New Species of Darkling Beetles from Central America with Systematic Notes (Coleoptera: Tenebrionidae)

ANNALES ZOOLOGICI (Warszawa), 2005, 55(4): 633-661

NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA WITH SYSTEMATIC NOTES (COLEOPTERA: TENEBRIONIDAE)

Julio Ferrer1 and Frode Ødegaard 2

1 Stora hundensgata 631, 13664 Haninge, Sweden, e-mail: [email protected] 2 Norwegian Institute of Nature Research, Tungasletta 2, N-7485 Trondheim, Norway, e-mail: [email protected]

Abstract.— A collection of Coleoptera Tenebrionidae from Central America has been studied and new species described and figured. The interest of this material principally consist in the method of sampling in the canopy and in the fact that for the first time the plant in which each specimen has been found was noted. Some systematic changes in the current classification of some genera, after Doyen and Tschinkel (1982) and Doyen et al. (1989) are introduced as results of morphological comparative study. Rhypasma Pascoe, 1871 is transferred to the tribe Stenosini from the Belopini. A total of 16 new species and one new genus from Panama are described and figured. Phymatestes agnei sp. nov., Rhypasma livae sp. nov., Lenkous ibisca sp. nov., Iccius monoceros sp. nov., Othryoneus triplehorni sp. nov., Paniasis kulzeri sp. nov., Gonospa similis sp. nov., Apsida simulatrix sp. nov., Brosimapsida gonospoides gen. and sp. nov., Epicalla elongata sp. nov., E. pygmaea sp. nov., E. aeneipes sp. nov., Strongylium vikenae sp. nov., Otocerus delicatus sp. nov. and O. angelicae sp. nov. The genus Paniasis Champion, 1886 is found to be identical to Pseudapsida Kulzer, 1961, created by monotypy for a species from Brazil: Paniasis brasiliensis (Kulzer, 1961) comb. nov. The systematic position of the genera Paratenetus Spinola, 1844, Rhypasma Pascoe, 1871, Calydonella Doyen, 1995, Othryoneus Champion, 1886, and Otocerus Mäklin, 1884 is commented.  Key words.— Coleoptera, Tenebrionidae, Central America, Panama, canopy.

Introduction accessed by two canopy cranes erected at the sites by the Smithsonian Tropical Research Institute (STRI). Besides A large part of the material presented in this study the Panamanian material, there are a few smaller collec- has been collected from two Panamanian tropical low- tions from Costa Rica, Nicaragua and Mexico. land forests. The first site, Parque Natural Metropolitano The canopy sampling was focused on Chrysomelidae (8°59´N–79°33´W, ca. 30 m a.s.l.), consist of 264 ha dry and Curculionoidea in order to estimate the host specif- tropical forest in Panama province, close to Panama City icity for phytophagous beetles (Ødegaard 2000a, 2003, and 2 km from the Pacific coast. The vegetation at this site Ødegaard et al. 2000) as a contribution to the debate is characterised by dominance of deciduous trees (30–35 around the magnitude of global arthropod species rich- m height) and lianas in the canopy. The other site is located ness (Ødegaard 2000b), that started as a consequence of in an evergreen, wet forest in San Lorenzo Protected Area Erwin´s controversial and inspiring assessments around (9°17´N–79°59´W, ca. 130 m a.s.l.) in Colón Province, the observation of a hyper-diverse beetle fauna occur- 4 km away from the Atlantic coast of the isthmus. This ring in a single tree species (Erwin 1982). Darkling forest includes 9,600 ha of relatively old-growth tropical beetles was regularly collected in these canopy samples forest of trees of 35–45 m height, and with lianas and and thus, the plant species in which the beetles were epiphytes occurring regularly in the canopy (Ødegaard recorded are available for most of species. in press). The beetles were collected by hand collecting Nearly 50% of species diversity in tropical forests is and by beating 10–40 m above ground. The canopy was probably restricted to the canopies (Ødegaard 2000b). This 634 J. FERRER and F. 0/DEGAARD percentage, however, is mainly based on phytophagous sions has been finished: revision of tribe Epitragini (Kulzer beetles, and it is unlikely that darkling beetles are hyper- 1964); genus Zopherus (Triplehorn 1972), today moved to diverse in the canopy based on observations during these own family Zopheridae; genus Doliema (Ardoin 1977), studies. On the other hand, as the fauna of darkling beetles and the genus Phaleria (Triplehorn and Watrous 1979) in the area is poorly known in general, it is not unexpected and genus Liodema (Triplehorn 1998). that several new species are to be found in this family. Conventional signs The study of Central American Tenebrionidae The systematic list of subfamilies, tribes and cur- The Central American components of the Family rent names follows Doyen and Tschinkel (1982) and Tenebrionidae sensu lato, including old Lagriidae and Doyen et al. (1989) with some commented exceptions. Allecullidae has been studied systematically only by Nomenclature is cited after Champion (1886), Gebien Champion (1886, 1913, 1917) with a classical work, the (1937–1939, 1940), Papp (1961). All specimens with- Biologia Centrali Americana, Heteromera, illustrating sev- out indication of collector name are collected by Frode eral species and permitting in many cases an immediate rec- Ødegaard. All described material is deposed in Frode ognition in visu of most species at generic level. Champion Ødegaard’s collection at NINA, the Norwegian Institute was the first specialist studying the genitalia of special dif- for Nature Research, Trondheim. Dubious taxa have ficult groups, as the old Cistelids (=Allecullidae). been compared with available type material or relevant A check list of Tenebrionidae and other Coleoptera authoritatively determined specimens, preserved in fol- of Mexico, West Indies, Central and South America has lowing Museums: Naturhistoriska riksmuseet, Stockholm been given by Blackwelder (1945). Another checklist of (F. W. Mäklin, 1884); Naturhistorisches Museum, Basel (H. America, north of the Panama canal was published by Gebien, H. Kulzer); The Natural History Museum, London Papp (1961). Listing 2483 taxa, this paper is an impor- (G. C. Champion, F. Bates), Muséum national d´Histoire tant contribution that gives the year and the page of the naturelle, Paris (M. Pic, P. Ardoin), Collection Julio Ferrer, original description, but in some cases (Mäklin 1884) Haninge (J. T. Doyen, C. Triplehorn), Synoptic collection the number of the page is wrong. of the Smithsonian Tropical Research Institute, Panama. Some factors explain current difficulties studying the Central American fauna: several numbers of taxa described by Colónel Casey and Maurice Pic are highly probably Taxonomy synonymies. For this reason and without examination of all available types of large genera such Platydema Laporte Phymatestes agnei sp. nov. de Castelnau (1831) or small species of Strongylium Kirby, (Figs 1–4, Photo 7) creation of new names to add the already very long list of obscure taxa, seems to be premature and hazardous. Type material. Holotype: ♂, Panama, Panama Prov., However, the access to available literature is not easy, Gamboa, 12.VI.1995. the books becoming very rare and exclusive; the types of Diagnosis. Length: 9.5 mm, maximum of width: 3.1 mm. many species are dispersed, preserved in many different The species of the genus Phymatestes has been recent- Museums or lost. For natural reasons, most American ly revised by Ferrer and Moragues (1998, 2000, 2001). specialists (Triplehorn 1965, Aalbu and Triplehorn 1985) This species belongs to the Phymatestes lamouri group. first initiated the study of American taxa north of Mexico, Characterised by moderate size, absence of tubercules on or focused on systematic problems of the higher classifica- elytra, unarmed legs and sub-parallel, elongate shape of tion of Tenebrionoidea (Doyen 1989, Doyen and Tschinkel body. This Phymatestes recall the habitus of Goniadera 1982, cladistic analyses of tribes and subfamilies) or most repanda (Fabricius), but differ by strong bicolour aspect, in nomenclatural problems (Spillman 1972, 1973). The pronotum blackish and dark, purpureous elytra and dif- German specialists studying the family (Freude 1967, ferent shaped aedeagus (cf. Fig. 4 and 5). 1968, Kulzer 1949, 1961, 1962, 1964) were concentrated Descriptiom. Shiny, glabrous, with some black, erect- to the South American representatives and the stud- ed hairs, only conspicuous laterally, sparsely disposed on ies of Central American taxa were rare and sporadic. tempora, pronotal sides and elytra. Marcuzzi (1953, 1991, 1994) studied the Caribbean and Head transverse, labrum rugose and pubescent, South American components, not the Central-American tegument irregularly and strongly punctured, epistome Tenebrionidae. Recently, Merkl and Maes (1991) pub- deeply impressed by a line, separating the raised supra- lished a current catalogue of Tenebrionidae for Nicaragua. antennal zones and the front. Eyes separated dorsally by The knowledge of the Central American Tenebrionidae is a distance superior to the diameter of an eye, measured still waiting contributions after Champion standard work, dorsally. Antennae long, reaching the middle of elytra, Biologia Centrali Americana. Only local study from the the third joint longer than the following, which are sub- vicinity of Jalisco, Mexico (Doyen 1988) and a few revi- equal, the ultimate two times longer as the preceding. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 635

Pronotum moderately transverse, 1.5 times as broad Managua, Las Flores, 5.II.1995, J. M. Maes (Museo as long, the anterior margin broadly opened, the anterior Entomológico de León, Nicaragua); Nicaragua, León, and posterior angles sub-right, the base bisinuate at each Carretera Poneloya-Chanchera, 29.I.1995, J. M. Maes and side, the sides strongly carinated, converging from before Collantes (coll. Julio Ferrer). the middle to the anterior board, constricted posteriorly. Description. Length: 5.8 mm, maximum of width: Tegument strongly and irregularly sculpted as head. The 1.8 mm. punctures here and there foveate and granuliferous. Elytra Black dull, pruinose. Head transverse, the epistome 2 times as long as broad, sub-parallel, the maximum of rounded, the genal zones long and round, becoming width after middle, rounded apically, depressed discally, sub-parallel before eyes. Eyes small and round, well strongly sculpted by irregular rows of foveate punctures, separated and placed on the posterior half of the head, the intervals moderately convex, shoulders marked by front moderately depressed, the tegument densely cov- humeral callus, the base straight, broader than pronotum. ered of round, shiny tubercules, concealing the cuticule. Legs simple, femora subclaviform, protibia sub-right, Antennae long, reaching the humeri, the third joint as the other subarcuate. long as the combined length of the two following joints, Etymology. Species named after Frode Ødegaard´s joints 4–8 sub-equal, quadrate, the following trans- son Agne. verse, the 10 shorter than the 9, the apical longer and rounded apically. Pronotum trapezoidal and strongly constricted back- Paratenetus foveithorax sp. nov. wards, the lateral zones broadly depressed, the anterior (Figs 20–24, Photo 5) board and the base straight; tegument densely covered of shiny tubercules as the head. Scutellum small, trans- Type material. Holotype: ♂, Panama, Panama Prov., versally rounded. Parque Natural Soberania, Plantation Road, 1.XII.2001. Elytra sub-parallel, about 2 times as long as broad, Description. Length: 3.3 mm, maximum of width: the base openly rounded, the shoulders not prominent, 1.5 mm. obtuse; with five finely carinate costae, included the lat- Castaneous brown, shiny, longely and sparsely cov- eral margin, the intervals flat and coarsely sculpted by ered with long yellowish hairs, normally winged. two rows of quadrate, foveate, contiguous punctures, Head microphthalmous, the epistoma broadly Ventral surface shiny, coarsely punctured. Mentum rounded, the antennal zones raised and convex, eyes very small, subtrapezoidal, finely punctured, the sub- small and round, very separated frontally. Tegument mentum large and transverse. Prosternal apophysis coarsely and sparsely punctured. Antennae pubescent, truncate; mesosternum sulcate by a fine medial longitu- the 6 first joints shiny and lighter than the following, the dinally traced carina, apophysis of metasternum broadly 7–8 black and shiny, the three ultimate blackish, duller. rounded, elytral epipleura finely and scarcely punctured Pronotum sub-cordiform, constricted basally, the and finely margined on the interior board. Abdomen sides irregularly contoured, without serrate denticula- shiny, apophysis of the first ventral sternite largely tion, the lateral margin broadly carinate and depressed, truncate, tegument of sternites strongly and densely disc with two callose elevations at each side. sculpted of tubercules, which are reclined and pointed Elytra ovate, acuminated apically, the base straight, backwards. The anal sternite unmargined. depassing the width of the pronotum, with callose Legs without diagnostic characters, the femora sub- posthumeral elevation, the disc moderately gibbose, clavate, the tibia straight. conspicuous laterally, epipleural zone very large, well Etymology. Species named after Frode Ødegaard´s separated from the lateral carina of the elytra by an daughter Liv. impression. Sculpture sparse, consisting in vestigial Remarks. The species of Rhypasma Pascoe (1871) rows of punctures, tegument longely and sparsely pilose, has been recently revised by Marcuzzi (1953, 1994, the hairs long and irregularly disposed here and there. 2002). The genus was transferred to the tribe Belopini Legs without diagnostic characters except the den- in the subfamily Lagriinae, by Doyen and Tschinkel tate protibia of the male. (1982). However, the characters of Rhypasma, indi- Etymology. Latin: foveithorax, alluding to the foveate cated close relationship with the tribe Stenosini, a group depressions of the pronotum. which was totally unrepresented between the genera used for cladistic analysis by Doyen and Tschinkel (loc. cit.). Rhypasma has to be retransferred to the Stenosini Rhypasma livae sp. nov. sensu Gebien (1937). The species of Rhypasma occur in (Figs 38–40, Photo 8) Haiti (1), in Venezuela (2), Trinidad (1), Bonaire (1), in the Amazonian region of Brazil (3), in Uruguay (1) and Type material. Holotype: ♀, Panama, Panama Prov., in Argentina (2). The genus and the species is new for Ancón, 16.II.2002, A. Ødegaard leg. Paratypes: Nicaragua, Central America. 636 J. FERRER and F. 0/DEGAARD

3 9 10 8 5 11 2 4

21 23

15 22 12 13 20 14

25 17 18 24 19

Figures 1–25. (1–4) Phymatestes agnei sp. nov. (1) Head and pronotum; (2) sculpture of elytra; (3) antenna; (4) aedeagus; (5) Goniadera repanda (F.); aedeagus. (6–11) Paratenetus tibialis Champion. (6) Aedeagus; (7) pronotum; (8) antenna; (9) protibia; (10) mesotibia; (11) metatibia. (12–13) Paratenetus grandicornis Motschoulsky. (12) Head and pronotum; (13) aedeagus. (14–17) Paratenetus ruficornis Champion. (14) Aedeagus; (15) protibia; (16) mesotibia; (17) head and pronotum. (18–19) Paratenetus sexdentatus Champion. (18) Aedeagus; (19) head and pronotum. (20–24) Paratenetus foveithorax. (20) Aedeagus; (21) antenna; (22) head and pronotum; (23) tibia; (24) profile; (25) Paratenetus denticulatus Champion; head and pronotum. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 637

Paniasis kulzeri sp. nov. lights). Paratypes: Panama, Colón Prov., San Lorenzo (Figs 41–46, Photo 6) Prot. Area, Flight intercepted trap, canopy, 21 m above forest floor, 25.V–4.VI.2004, R. Didham et al.; ídem, Type material. Holotype: ♂, Panama, Panama Prov., 1–13.X.2003, R. Didham and L. Fagan leg. Museum of Gamboa, 10-V.1995. Natural History, Budapest, ♂. ♀. Description. Length: 6.2 mm, maximum of width: Description. Length: 2.8 mm, maximum of width: 2.8 mm. 0.7 mm. Glabrous, dull, the elytra with a silky aspect, brown- Light castaneous, glabrous, somewhat translucidous, ish with reddish buccal appendages, legs and four basal, normally winged. Very similar in shape to Sicinus shiny antennal joints, the following joints darker and gutemalensis Champion, 1886, but clearly different by duller. Normally winged. proportionally more elongate body, not so broad, sub- Epistoma rounded, supra-antennal zones moderately quadrate shape of pronotum, shorter elytra and cephalic raised and convex, the eyes reniform, separated frontally by configuration, which in this new species lack the four a distance equivalent to the diameter of an eye, measured characteristic tubercles of this species, the transversal dorsally; tegument densely covered of punctures, separated callus on the vertex and the compact and dorsally sul- by a distance equivalent to the diameter of a puncture. cate mandibles. Antennae long, reaching the middle of elytra, the third This uniformly coloured species also seems to be joint as long as the fourth, the following becoming closely related to Iccius cylindricus Champion, 1886: 148, longer and progressively punctured. which is bicoloured, ornated by reddish mandibles, cas- Pronotum trapezoidal, the anterior board and the taneous legs and elytra, except apical third, with black base bisinuate, sides feebly constricted anteriorly, sub- head, antennae, pronotum and apex of the elytra, and parallel backwards, anterior angles sub-right, posterior presenting another cephalic sculpture, consisting in two obtuse, base unmargined, lateral carina finely traced small tubercules longitudinally disposed frontally. from sides to anterior margin of pronotum. Tegument Kulzer (1949) has described two new species of this sculpted as head, the punctures fine, rounded, here and genus: I. elongatus from Costa Rica and I. brevipennis, there contiguous, generally well separated. from Brazil. I. elongatus is darker, olive-green and the Elytra sub-parallel, ovate apically, about 1.5 times as head is bicorne like in brevipennis. This new species is long as broad, the maximum of width at middle, Callus the only species of this genus orned by a single horn humeral distinct, lateral carina perfectly visible dorsally, at middle of front. A more detailed description of this sculpted with regular rows, finely incised with small, species has to be given in a revision of Gnatocerus and round, separate punctures, becoming superficial api- related genus. cally, intervals flattened, moderately convex on sides, Etymology. Greek: µοηοκεροσ: with a single horn, finely and sparsely punctured. alluding the peculiar cephalic configuration of this insect. Legs slender with characteristic dilated protarsi. Etymology. Species named after Hans Kulzer, the late specialist of Tenebrionidae. Othryoneus triplehorni sp. nov. Remarks. The genus Pseudapsida Kulzer, 1961, separate (Figs 74–76, Photo 1) from Platydema by strongly dilated protarsi, was created by monotypy by Kulzer (1961), to receive a 12 mm long new Type material. Holotype: ♂, Panama, Colón Prov., species from Brazil: P. brasiliensis. This second species from San Lorenzo, Prot. Area, 11.V.2004. Paratype: ♀, Panama, Panama is very similar in facies, but completely different Panama Prov., Gamboa, 13.VI.1995. by much smaller size, unmargined pronotum, coarser pro- Diagnosis. Holotype – length: 5.1 mm, maximum of notal punctures and uniform intervals of elytra. width: 3 mm. Paratype – length: 6.5 mm, maximum of The genus Paniasis, created by Champion 1886: 206, width: 3.5 mm. to receive a single species from Mexico, is a senior syn- Similar in shape to a Platydema, but lacking the cari- onyme of Pseudapsida Kulzer, 1961 syn. nov., created nate surface of legs, characteristic of Diaperini. by monotypy, for a species from Brazil, characterized by Description. Body glabrous, head and pronotum, strongly dilated protibia. We recommend the transfer legs and the 5 basal joints of antennae, reddish orange, of this species to the genus of Champion: proposing the following black and duller; elytra orange with four Paniasis brasiliensis (Kulzer, 1961) comb. nov. discal black patches. Normally winged. Head strongly transverse, microphthalmic, antennae Iccius monoceros sp. nov. short, not reaching the base of pronotum, the third joint (Figs 48–49) two times as long as broad, as long as the fourth, the following 4–6 joints subquadrate, the following becoming Type material. Holotype: ♂, Panama, Colón Prov., progressively dilated, the apical joint round. Eyes round, San Lorenzo Prot. Area, 13.X.2003, R. Kitching leg. (at small and well separated, the epistoma truncate, the 638 J. FERRER and F. 0/DEGAARD

35 27

30 31

33 28

34 43

44 41 46

42 40 45

Figures 26–46. (26–27) Paratenetus sp. (26) Aedeagus; (27) head and pronotum. (28–29) Paratenetus constrictus Champion. (28) Aedeagus; (29) head and pronotum; (30) Paratenetus sp. Habitus. (31–32) Paratenetus sp. cf. crenulatus Champion. (31) Head and pronotum; (32) elytral sculpture. (33–37) Lenkous ibisca sp. nov. (33) Habitus; (34) antenna; (35) prosternum; (36) meso- and metasternum; (37) aedeagus. (38–40) Rhypasma livae sp. nov. (38) Ovipositor; (39) antenna; (40) habitus. (41–46) Paniasis kulzeri sp. nov. (41) Habitus; (42) aedeagus; (43) antenna; (44) protarsi; (45) mesotarsi; (46) metatarsi. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 639 labrum well separated by a cireous membrane, sparsely, L., black with a brownish tint, with reddish, testaceous yellowish pubescent. antennae, buccal appendages and legs. Pronotum transverse, two times as broad as long, Head rounded, the eyes separated at front by a the anterior board and the base bisinuate, the anterior distance equivalent to 1.5 times the diameter of an angles obtuse, the posterior sub-right, the sides regu- eye, measured dorsally, finely and sparsely punctured. larly convergent from base to anterior board. Tegument Antennae with the third joint moderately long, the strongly and densely covered of elongate punctures. fourth shorter, a little longer than broad, the following Elytra ovate, 1.5 times as long as broad, with strongly becoming shorter and progressively dilated, the apical carinate shoulders and broadly semicircular scutellum. joint rounded. Each elytron maculate by a long irregular black patch Pronotum large, and short, strongly transverse, 2 occupying the 3–4 discal and 5–6 lateral intervals, at the times as broad as long, the anterior board broadly first elytral third and two other transversally disposed opened, with obtuse anterior and posterior angles. The and smaller in the posterior third. lateral carina well traced into the lateral portion of the Mentum rounded and finely punctured, contrasting anterior board and the base, which is impressed at each with the extremely strongly and densely rugose punc- side. Tegument sparsely and finely punctured. Scutellum tures of the subgular zone. Gular zona moderately glo- small and triangular, depressed between elytra. bose and punctured, unpunctured at middle. Propleura Elytra semi-sphaeric, globose, largely margined lat- strongly punctured. Apophysis of prosternum finely erally, the base completely adapted to the shape of the reborded, rounded apically. Mesosternum roundly exca- base of pronotum, permitting hermetic closing of body, vate, coarsely rugose punctate at each side, separated shoulders obtuse, convex discally, with fine, but con- from metasternum by a stroge carina. Espisternal zones spicuous rows of well separated punctures. strongly punctured, as the elytral epipleura. Apophysis Propleura lise and broadly excavate at each side to of first ventral sternite margined, triangulary rounded. receive the anterior legs, prosternal apophysis reclined Sternites I–III lateraly margined and punctured, IV and and strongly sulcated and margined. V with membrana, obsolescently punctured and unmar- Mesosternon very short, longitudinally carinate at gined. Legs simple, without particular features, lacking middle and sculpted with two diagonally disposed at lateral carina. middle convergent carine at each side. Metasternon Etymology. Species named after Charles Triplehorn, opened in “U” at middle, the episternes dull, coarse- eminent specialist of Tenebrionidae. ly and scarcely punctured, the discal zone rounded, Remarks. The habitus of the female recall the shape of strongly and densely punctured. a Platydema. The pronotum of the male holotype is more Etymology. IBISCA, acronyme of the project: trapezoidal, not so large and strongly punctured (Fig. 74). Investigating the BIodiversity of Soil and Canopy The species of Othryoneus (Champion, 1886: 245) Arthropods. has been placed in the old tribe Cnodalonini, formerly Remarks. Lenkous Kaszab, 1973 is a myrmecophilous in the subfamily Coelometopinae. Before the descrip- genus from Brazil described by monotypy to receive a tion of this new species the types of all species described species described by him at this occasion. Kaszab placed by Champion (loc. cit.) and Kulzer (1964) have been it in the tribe Cnodalonini after Gonospa. The systematic examined. A revision of this genus is under progress. position of this genus, apparently similar in facies to the This new species, one of the smallest of the genus, has to genera of the tribe Leiochrini and of the Nilioninae (sensu be placed after O. serrivittatus Kulzer, 1964. Doyen et al. 1989) is preliminary. Provisionally, we think that the place of this new genus is near the Leiochrini, probably exiging the creation of a new tribe. The genus Lenkous ibisca sp. nov. and species are new for Central America. (Figs 33–37, Photo 16)

Type material. Holotype: ♂, Panama, Colón Prov., Gonospa similis sp. nov. San Lorenzo Prot. Area, extracted by Berlese-Tullgren (Figs 60–62, Photo 13) from epiphytes collected in the canopy, 27.IX.2003, N. Winchester and K. Jordan leg. Paratypes: (sex not Type material. Holotype: ♀, Panama, Panama Prov., examined) Panama, Colón Prov. San Lorenzo Prot. Parque Natural Metropolitano, 24.IV.1995 (on Bonamia Area, 8.X.2004, J. Bail leg. (canopy fogging) Museum of marapoides). Paratype: ♀, Panama, Panama Prov., Parque Natural History, Budapest and coll. J. Ferrer. Natural Metropolitano, 2.II.1995 (on Dioclea guianensis); Description. Length: 2.2 mm, maximum of width: ídem, 15.I.1996; ídem: 5–6.IV.2004; San Lorenzo Forest, 1.3 mm. IBISCA, Teneb65, Museum of Natural History, Budapest. Body shiny, glabrous, normally winged, completely Description. Length: 2.5 mm, maximum of width: rounded and convex, recalling the facies of a Coccinella 1.5 mm. 640 J. FERRER and F. 0/DEGAARD

Glabrous, dark brownish aeneous, with a transluci- Etymology. Latin: allusion to the similarity of this dous red-brownish tint, with antennae, buccal append- species with Gonospa. ages and legs brown. Normally winged. Remarks. This species is another case of conver- Head hidden, concealed by pronotum, epistome gence, recalling the habitus of Gonospa phaedonoides. broad, straight, truncate, well separated from the front However, Gonospa are easily recognised by 6-joints dark by a transversal, well impressed line. Antennae slender club and massive metallic dark legs. This species differ reaching the shoulders, the club formed by 5 moderately by progressively incrassated antennae and very gracile, progressively dilated joints. reddish light legs. Using Kulzer (1961: 218) key, this Pronotum strongly transverse, broadly opened ante- new species can be placed between A. bucardi Bates and riorly, about 2 times as long as broad in semi-circle, A. cubaensis Kulzer (loc. cit.), using Triplehorn (1969), lateral margins finely carinate, anterior angles broadly between bucardi and terebrans Champion. obtuse, sides regularly curved, posterior angles sub- right, base unmargined and bisinuate at each side; tegument finely, sparsely and superficially punctured, finely Brosimapsida gen. nov. sculpted by an isodiametrical web of transversal micro- (Figs 69–73, Photo 14) scopic rides, only perceptible by high magnification, the prebasal zone largely depressed. Diagnosis. A new genus is necessary for the recep- Elytra globose, becoming ovate apically, about 1.5 tion of a remarkable species, which will be recognized times as long as broad, lateral carina strong and hardly by strongly shiny, aeneous, glabrous, semi-hemisphaeric visible dorsally, forming with the lateral intervals a form as other species of Apsida, combined by the progres- broadly depressed apical zone. sively dilated, asymetrical, subserrate antennae (Fig. 69) Etymology. Allusion to the similar aspect of this spe- and the peculiar obtusely serrate protibia (Fig. 71). cies with Gonospa phaedonoides Champion. Type species. Brosimapsida gonospoides sp. nov. (by monotypy); gender feminine.

Apsida simulatrix sp. nov. (Figs 63–65, Photo 15) Brosimapsida gonospoides sp. nov.

Type material. Holotype: ♀, Panama, Colón Prov., Type material. Holotype: ♀, Panama, Colón Prov., San Lorenzo Prot. Area, 4.V.2001 (on Virola elongata). San Lorenzo Prot. Area, 21.XII.2001 (on Brosimum Paratypes: Panama, Colón Prov., San Lorenzo Prot. utile). Paratype: ♂, Panama, Colón Prov., San Lorenzo Area, 3.IX.2003, H. Barrios and F. Ødegaard leg.; ídem, Prot. Area, IBISCA, flight-intercept trap, 35 meters 11.V.2002 (on Arrabidaea verrucosa); ídem, 30.XI.2001, above ground, 1–17.X.2003, R. Didham and L. Fagan (on Marila laxifolia); ídem, 24.I.2002. leg. (Museum of Natural History, Budapest). Description. Length: 2.5 mm, maximum of width: Description. Length: 3.3 mm, maximum of width: 1,8 mm. 2.5 mm. Shiny, glabrous, aeneous, with a strong, translucidous, Body semi-sphaeric, glabrous, shining aeneous, with reddish brown tint, with antennae, buccal appendages buccal appendages, three basal joints of antennae and and legs, yellowish reddish, more or less darker. Normally legs reddish brown, ventral surface shiny brown, with winged. lighter epipleura. Normally winged. Head truncate anteriorly, hidden by the board of Head transverse, epistome truncate, genae sinuate pronotum, epistome truncate, separated from the front and curved backward, the supra-antennal zone deeply by a transversal line, eyes well separated frontally by foveate, eyes transverse, separated frontally by a distance a distance superior to 3 times the diameter of an eye, equivalent to 1.5 times the diameter of an eye, measured measured dorsally. dorsally; tegument finely, irregularly punctured. Pronotum discally convex, anterior board broad- Pronotum broadly transverse, 2 times as broad as ly opened, anterior angles broadly obtuse, posterior long, broadly opened anteriorly and curved basally, angles sub-right, sides regularly curved, base unmar- anterior angles broadly obtuse, posterior angles sub- gined, bisinuate at each side, largely and transversally right, lateral sides regularly curved. Tegument finely and depressed; tegument extremely fine, sparsely and super- irregularly punctured, as the head. ficially punctured, finely sculpted by an extremely fine Elytra globose and semi-sphaeric, shoulders broadly isodiametrical somewhat transversal reticulation, only obtuse, elytral carina invisible dorsally. perceptible by high magnification. Gular zone strongly convex, separated by a fine, Elytra ovate, globose; shoulders rounded, lateral impressed line from submentum, maxillar palpi dilated carina invisible dorsally, forming apically with the lateral and securiform, the marginal board well separated and interval a broad depressed zone. lighter, of a cireous aspect. Propleura excavate to receive NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 641

47 51 50

55 54

52 53 56 57 49

68 48 61 69 58

65 59

67 63

62 64 60

73 76

75 72 71 70 74

Figures 47–76. (47) Platydema sp. of sobrinum-group. Aedeagus. (48–49) Iccius monoceros sp. nov. (48) Aedeagus; (49) head and pronotum. (50–53) Ca- lymmus cucullatus Pascoe. (50) Pronotum ♀; (51) ídem ♂; (52) ovipositor; (53) aedeagus. (54) Ozolais elongata Champion. Pronotum ♀. (55–59) Antima- chus coriaceus Lacordaire. (55) Protibia; (56–57) cephalic horn; (58–59) aedeagus. (60–62) Gonospa similis sp. nov. (60) Aedeagus; (61) antenna; (62) pronotum. (63–65) Apsida simulatrix sp. nov. (63) Antenna; (64) pronotum; (65) ovipositor. (66–68) Gonospa phaedonoides Champion. (66) Pronotum; (67) antenna; (68) ovipositor. (69–73) Brosimapsida gonospoides gen. and spec. nov. (69) Antenna; (70) habitus; (71) protibia; (72) ovipositor; (73)apopysis of mesosternum. (74–76) Othryoneus triplehorni sp. nov. (74) Habitus; (75) aedeagus; (76) antenna. 642 J. FERRER and F. 0/DEGAARD the legs. Apophysis of prosternum recurved between Etymology. Aeneipes: latin alluding the cupreous coxae, broad, pointed apically. Mesosternon short, with aspect of legs of this insect. a small open apophysis to receive the prosternal piece (Fig. 73), which is subrectangular, not U or V-shaped, as normal. Metasternum unpunctured, epipleura excavate Epicalla elongata sp. nov. to receive the legs. (Figs 77–78, 86, Photo 9) Etymology. Composite name: alluding to the closely related genus and to the tree species Brosimum utile in Type material. Holotype: ♂, Panama, Colón Prov., which this insect was found. San Lorenzo Prot. Area, 11.V.2001. (on Dendropanax arboreus). Description. Length: 12 mm, maximum of width: Epicalla aenipes sp. nov. 3.5 mm. (Fig. 84, Photo 10) Epistoma truncate, supra-antennal zones raised in rapport to the level of the epistoma, but lower than the Type material. Holotype: ♀, Panama, Panama Prov., front, separated by a transversal impression, eyes large, Parque Natural Metropolitano, 17.IV.1995. transverse, subglobose, separated frontally by a distance Paratype: ♂, Panama, Colón Prov., San Lorenzo equivalent to the combined length of the two first anten- Forest, IBISCA, 12.V.2004 (Museum of Natural History, nal joints, tegument irregularly punctured. Antennae Budapest). reaching the shoulders, the first joints shiny, the follow- Description. Length: 10.0 mm, maximum of width: ing darker and duller, coarsely punctured. 4.2 mm. Pronotum short, 1.5 times as long as broad, the ante- Body shining metallic green, glabrous, aeneous, rior board straight, the base a little bisinuate at each side, with a cupreous tint on legs, antennae black and dull. anterior angles broadly rounded, posterior angles sinuate, Head transverse, macrophthalmic, the eyes oblongly just before the base and acute, strongly carinated later- globular, reniform on profile, separated by a distance ally, anteriorly and basally, with a prebasal impression at equivalent to half diameter of the eye, measured dorsally. each side, united by a prebasal, transverse impression. Epistome convex, truncate; labrum as short as the buccal Tegument irregularly punctured. Scutellum very reduced. membrana and pubescent; supra-antennal zones raised; Elytra elongate, two times as long as broad, sub-par- canthus ocular rounded; front separated from the epis- allel in shape, rounded apically, the base broader than toma by a transversal line, deeply depressed between eyes; the width of the pronotum basally, the lateral carina tegument strongly, irregularly and sparsely punctured. strong and entirely visible dorsally, except at apex, rows Antennae moderately long, reaching the shoulders. strongly lineate, the scutellar striae strongly depressed Pronotum transverse, about 1.5 times as broad as forming a long postscutellar zone, intervals moderately long, anterior board convex, strongly margined at each convex and sparsely punctured. side, anteriorly and basally, anterior angles broadly Legs without diagnostic characters, except the mes- rounded, posterior angles sub-right, sides regularly ofemora, which are orned by a short fringe of yellowish curved forwards, abruptly constricted and sinuate basal- hairs ventrally. ly; base strongly bisinuate at each side; tegument strongly Etymology. Allusion to the peculiar elongate form and irregularly punctured, the punctures microgranifer- of this species. ous, here and there confluent, generally well separated; basal third with three impressions united by a transversal impressed zone. Epicalla pygmaea sp. nov. Elytra 2 times as long as broad, the maximum of (Fig. 85, Photo 12) width after middle, sub-parallel becoming enlarged, ovate; shoulders prominent, depassing the width of Type material. Holotype: ♂, Panama, Panama Prov., pronotum, basal zone depressed, tumefact, elytral carina Parque Natural Metropolitano, 4.VI.1995 (on Luehea strongly visible humerally, constricted apically; 8th inter- seemannii). Paratype: ♀, ídem, 1.V.1995 (on Luehea see- val finely carinate apically 5 discal intervals strongly mannii); ídem, 5–6.V.2004. lineate, the punctures only conspicuous laterally; lateral Description. Length: 5.8 mm, maximum of width: intervals strongly sulcate and punctured. 2.5 mm. Ventral zone shiny, glabrous, apophysis of prosternum Body shining strongly metallic, dark, aeneous, with strongly reborded and pointed, the apex however, obtuse- reddish cupreous disc on elytra. Normally winged. ly rounded. Mesosternon short, concave; metasternon Head oblong, strongly transverse, epistoma truncate, shiny and convex, longitudinally impressed at middle, straight and deeply transversally depressed, supra-anten- extremely fine and sparsely punctured at middle; prop- nal zones deeply and transversally impressed and foveate leural and epipleural zones strongly punctured. before eyes, which are globose, separated frontally by a dis- NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 643

77 79 80 81

84 85

86 87

83 88 89

91 96 101 102

104

103

95 92 94

90 93 97 98 99 100

Figures 77–104. (77–78, 86). Epicalla elongata sp. nov. (77) Aedeagus; (78) protarsi; (86) habitus. (79) Epicalla sp. Aedeagus. (80) Calydonella lisa Doyen. Aedeagus. (81–83, 87) Epicalla instriata Pic. (81) Aedeagus; (82) protarsi; (83) antenna; (87) habitus. (84) Epicalla aeneipes sp. nov. Habitus. (85) Epicalla pygmaea sp. nov. Habitus. (88–90) Strongylium vikenae sp. nov. (88) Sculpture of pronotum; (89) sculpture of elytra; (90) aedeagus. (91–92, 99–100) Otocerus delicatus sp. nov. (91) Aedeagus; (92) antenna; (99) sculpture of pronotum; (100) sculpture of elytra. (93–95, 97–98) Otocerus angelicae sp. nov. (93) Antenna; (94) protibia; (95) aedeagus; (97) sculpture of pronotum; (98) sculpture of elytra. (96) Otocerus flavipennis Mäklin. Aedeagus. (101–102) Poecilesthus variipes Champion. Aedeagus. (103) Strongylium? clavicornis Champion. Habitus. (104) Strongylium? sp. aff. clavicornis Champion. Habitus. 644 J. FERRER and F. 0/DEGAARD tance superior to the diameter of an eye, measured dorsally; Elytra depassing the width of the pronotum, deeply front deeply impressed between eyes, tegument finely and sulcate of darker striae, consisting of dense rows of black sparsely punctured. Antennae short, reaching the humeri, punctures, the alternate intervals larger than the others. the third joint short, as long as the fourth but smaller, the Ventral surface metallic green, shiny, strongly punc- following becoming progressively transverse and sub- tured, glabrous, except the apex of the anal sterni- equal, the apical joint longer and rounded apically. te. Mentum small and finely punctured, pubescent. Pronotum transverse, two times as broad as long, Maxillar palpi truncate and orned by a margin of cireous trapezoidal, the sides rounded anteriorly and sub-paral- aspect apically; submentum subpentagonal and concave lel backwards, the margin deeply carinated, the anterior the pregular zone strongly sulcate, strongly rugose and and basal board strongly margined, the anterior angles transversally sulcate at middle. Gula convex and shiny. rounded, the posterior sub-right, the base bisinuate and Eyes separated ventrally by a distance equivalent to 0.7 with a little foveate impression at each side; tegument times the diameter of an eye, measured ventrally. Board finely, superficially and sparsely punctured. anterior pf pronotum finelly margined and orned by a Elytra broader than the base of pronotum, strongly fringe of brownish-yellow hairs. coarsely and densely dilated after middle, about 1.5 times as long as broad, punctate, the punctures nearly contiguous. Apophysis the shoulders prominent, strongly carinated, the lateral of prosternum large, rounded apically and recurved carina entirely visible dorsally, elytral rows lineate, the between coxae, but visible laterally. punctures obsolescent discally, the seventh interval Mesosternon coarsely punctured, anteriorly and lateral- finely carinated backwards and rounded apically. ly, deeply excavated at middle, between coxae and transver- Etymology. Pygmaea, latin: small, allusion to the sally rugose. Metasternon anteriorly truncate and forming reduced dimensions of this species. with the posterior zone of the mesosternum a deep excavate depression, probably to conserve humidity. Complete evaporation of water conserved in this natural recipient Strongylium vikenae sp. nov. was obtained only after 35 minutes in a room temperature (Figs 88–90, Photo 2) of 21°C. Epimeral zone densely and coarsely punctured, the punctures becoming sparse, smaller and feebly transverse Type material. Holotype ♂.Panama, Panama Prov., at middle, forming a combination of superficial poinst and Barro Colorado Isl. 11–13.V.2004. Paratypes: 2 ♀♀: transverse rides, becoming obsolescent backwards; disc Panama Prov. San Lorenzo prot. Area, 9–28.V.2005, finely impressed longitudinally at middle, the impression G. Curletti leg. (2 specimens in Museo Civico Storia becoming larger and excavated basally. Elytral episternes Naturale, Carmagnola and coll. J. Ferrer, Haninge); same finely margined and rugose apically. location, 22–31.V.2004 (one female collected at light). Abdomen shiny, irregular, sparsely and strongly Diagnosis. This beautiful species recalling the habi- punctured, the punctures well separated but sometimes tus of Otocerus flavipennis Mäklin, 1884, from Brazil, is confluent or contiguous, becoming almost isolate at one of the largest American Strongylium. However, the middle of each sternite. Ventral apophysis lanceolate shape of the antennae and the aedeagus (cf. Figs 90 and and strongly margined. Sides of sternites 1.2 and 4 96) combined by other external characters, clearly indi- depressed at each side. The anal sternite small, unmar- cate the convergence of both species. Morphologically, gined, depressed and finely pubescent apically. this new species is likewise similar to Strongylium car- Etymology. Species named after Åslaug Viken, the inipenne Champion, 1888: 374, which is much smaller wife of Frode Ødegaard. and exhibiting a dark patch posterior on elytra. Description. Length 31 mm, maximum of width: 5 mm. Body metallic green with yellowish, shiny elytra. Otocerus angelicae sp. nov. Normally winged. (Figs 93–94, 97–98, Photo 4) Head coarsely punctured, the margin of the clypeus curved, front deeply foveate between the eyes, which are Type material. Holotype: ♂, Panama, Panama Prov., separated frontally by a distance equivalent to 0.5 times Barro Colorado Isl., 11–13.V.2004. the diameter of an eye, measured dorsally. Antennae long, Description. Length: 14 mm, maximum of width: 3 mm. reaching the anterior third of the elytra, the third joint Body glabrous, metallic green, moderatelly shiny, sub-equal to the following, shiny and finely punctured, the with dull, black antennae. Normally winged. following darker and dull, covered of short black setae. Head strongly macrophthalmic, the board of the Pronotum feebly transverse, nearly quadrate and epistome rounded and punctured, the genal, supra- subcylindric, without lateral protuberances, strongly -atennal zones strongly raised, brownish. Eyes large margined basally and anteriorly, the carina becoming and nearly contiguous, separated frontally by a distance larger at middle of the anterior board. Lateral zones equivalent to an ocular facette, the separation back- unmargined, covered of coarse punctures as the head. wards equivalent to the length of the first antennal joint. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 645

2 3

Photos 1–7. (1) Othryoneus triplehorni sp. nov., female; (2) Strongylium vikenae sp. nov.; (3) Otocerus delicatus sp. nov.; (4) Otocerus angelicae sp. nov.; (5) Paratenetus foveithorax sp. nov.; (6) Paniasis kulzeri sp. nov.; (7) Phymatestes agnei sp. nov. 646 J. FERRER and F. 0/DEGAARD

8 12 9

14 11

15 16

Photos 8–16. (8) Rhypasma livae sp. nov.; (9) Epicalla elongata sp. nov.; (10) Epicalla aeneipes sp. nov.; (11) Epicalla instriata Pic; (12) Epicalla pygmaea sp. nov.; (13) Gonospa similis sp. nov.; (14) Brosimapsida gonospoides gen. and sp. nov.; (15) Apsida simulatrix sp. nov.; (16) Lenkous ibisca sp. nov. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 647

21 20

Photos 17–21. (17) Strongylium sp. 1; (18) Strongylium sp. 2; (19) Strongylium sp. 3; (20) Strongylium sp. 4; (21) Strongylium? sp. aff. clavicornis Champion. 648 J. FERRER and F. 0/DEGAARD

Vertex forming a sharply triangular zone between eyes, Epistoma truncate, labrum darker, brown, separated tempora constricted, Antennae very long and slender, by a transversal membrane, maxillar palpi moderately reaching the apical third of elytra, the first joint as short dilated apically; head macrophthalmic, the eyes sepa- as the third, which is much shorter than the following (a rated frontally by a distance inferior to the width of the diagnostic character of gen. Otocerus). Joints 4–10 sub- first antennal joint, vertex pointly triangular, between equal, duller, finely dilated at the external apex, the last eyes, tegument coarsely punctured, finer anteriorly, joint elongate and rounded apically. supra-antennal zones raised, the ocular canthon short, Pronotum sub-cylindrical, nearly trapezoidal, the occupying about one third of the width of the eye, base a little larger than the anterior board, with a measured laterally; eyes strongly reniform, globose in large, basal impression at middle and two at each side. dorsal view. Antennae long, reaching the apical third of Tegument coarsely punctured, each point microgranif- elytra, the third joint aboint ½ the length of the follow- erous, irregularly disposed, some times contiguous, the ing (diagnostic character of s. gen. Otocerus), the joints tegument isodiametrically reticulate. Scutellum round 4–9 sub-equal, but becoming progressively dilated, the and very finely punctured. 9 and10 a little shorter, the last joint becoming elongate Elytra elongate, three times as long as broad, the and rounded apically. shoulders rounded, acuminate apically, with dehiscent Pronotum sub-cylindrical, the sides becoming a little apex, sulcate by nine striae, included the scutellar, con- convergent to the anterior board, sub-parallel and orned vergent two and two basally and consisting in well incised with a little lateral protuberance after middle, irregularly rows of sub-quadrate, sub-foveate points, united by a coarsely and strongly punctured. continuous line. Legs very long and slender, the protibia Elytra shining metallic green, with a brownish tint, shorter and curved, the meso and metatibia straight, the sub-parallel, acuminated apically, coarsely sulcate by tarsi very long, without diagnostic characters. densely punctured rows, the intervals convex and finely Etymology. Species named after Julio Ferrer’s daugh- punctured. ter Angelica. Legs very slender and long, without diagnostic char- This species is closely similar to Otocerus attenuatus acters. Champion. The following table permitting the separa- Etymology. Delicatus, latin: gracious, alluding to the tion of both species: gracility of this species. Remarks. Otocerus has been separated from O. attenuatus Champion O. angelicae sp. nov. Strongylium by third antennal joint reduced, and con- eyes widely separated frontally eyes nearly contiguous stantly shorter than following. However, dimorphical front finely punctured front strongly punctured legs occur in some species of Otocerus. joints 4 and following violaceous joints 4 and following dull, black pronotal protuberance pronotal lateral protuberance unconspicuous conspicuous apex of elytra pointed apex of elytra dehiscent elytral punctures oblong elytral punctures round to Annotated check-list of species subquadrate Intervals flat to beyond the middle intervals convex Subfamily Lagriinae base of tibia and tarsi reddish legs uniformely black (except tribe Statirini, which will be studied in a sepa- length: 11 mm length: 14 mm rate paper)

Tribe Lagriini Otocerus delicatus sp. nov. Lagria villosa Fabricius, 1781 has been recently intro- (Figs 91–92, 99–100, Photo 3) duced in Brazil (Paraná, Foz de Iguazu, 21.VIII.2000, Sta Catherina, Florianopolis, Praia do Colón Prov. Santinho, Type material. Holotype: ♂. Panama, Panama Prov., 24.VIII.2000) and Argentina (Corrientes, Cataratas, Parque Natural Metropolitano, 5–6.V.2004. Paratype: ♂. 1.IX.2000). The presence of this African Lagria in Panama, Colón Prov., San Lorenzo Prot. Area, 9.V.2001. Central America is probably a matter of time. Description. Length: 7.5 mm, maximum of width: 1.8 mm. Tribe Lupropini Glabrous, metallic green, shiny, with a castaneous to Currently, North American specialists after Doyen reddish-brown tint, head, legs, anterior cephalic zone, et al. (1989) ignored the tribe Lupropini sensu Ardoin buccal appendages and three first basal antennal joints, (1961), including these insects in the tribe of Goniaderini ventral zone of body, anterior and basal carina boarding or other tribal groups. The genus Paratenetus Spinola the pronotum, lateral carina of elytra, strongly shiny (1844: 116) has been transferred by Doyen et al. (1989) to cataneous-brown. Antennae black, with two apical joint the tribe Belopini. However this genus is closely related lightly yellowish. Normally winged. to the African genus Enicmosoma (sensu Ardoin 1959), NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 649 and belongs to Lupropini, sensu Ardoin (1959, 1961) in Paratenetus constrictus Champion, 1892 all essential morphological characters. None adequate (Figs 28–29) keys exists and the genus need a revision. Costa Rica, Guanacaste Prov., Sta Elena, 10.II.1998; Paratenetus denticulatus Champion, 1886 Panama, Panama Prov., Parque Natural Metropolitano, (Fig. 25) 18.III.1995; (on Enterolobium cyclocarpum) (3).

Panama, Colón Prov., San Lorenzo Prot. Area, Paratenetus sp. cf. crenulatus Champion, 1886 9.V.2001 (on Nectandra purpurascens); ídem, 20.IX– (Figs 31–32) 30.X.2003; (on Inga pezizifera); ídem, 27.X.2001 (on Inga cocleensis); Panama Prov., Parque Natural Metropolitano, Panama, Colón Prov., San Lorenzo Prot. Area, 19.I.1996 (on Luehea seemannii). 12.V.2004 (on dead branches in the understorey). A species, characterised by serriate rows of elytral Paratenetus tuberculatus Champion, 1886 punctures (Fig. 32).

Panama, Panama Prov., Parque Natural Metropolita- Paratenetus nigricornis Champion, 1886 no, 17. IV. 1995 (on Combretum fruticosum). Closely related to a syntype of P. tibialis Champion, Panama, Colón Prov., San Lorenzo Prot. Area, 1886 (Figs 6–11), but smaller. 24.V.2001; ídem, 12.IV.2002 (on Tocoyena pittieri); ídem, 24.III.2004 (on Tocoyena pittieri, on dead branches in Paratenetus ruficornis Champion, 1886 the canopy). (Figs 14–17) Paratenetus foveithorax sp. nov. Panama, Panamá Prov., Cerro Campana, 650 m a.s.l., (Figs 20–24, Photo 5) 16.V.1996. Panama, Panama Prov., Parque Natural Soberania, Paratenetus sexdentatus Champion, 1886 Plantation Road, 1.XII.2001. (Figs 18–19) Paratenetus sp. Panama, Chiriqui Prov., Palo Alto, 1400–1600 m (Figs 26–27) a.s.l. 10.VIII.1995; Colón Prov., San Lorenzo Prot. Area, 27.VII.2001 (on Nectandra purpuracens); ídem, 3.V.2001 Panama, Colón Prov. San Lorenzo Prot. Area, (on Cydista aequinoctialis); ídem, 2.VI.2001, (on Clusia I.X.2003 (on Terminalia amazonica). minor); ídem, 15.V.2003 (on Tocoyena pittieri); ídem, 5.X.2003, (on dead branches of Terminalia amazonica Paratenetus sp. in the canopy). (Fig. 30) Remark. This species is closely related to Paratenetus auritus Mäklin (1875), from Amazonas (Brazil, Cabessa Panama, Panama Prov., Parque Natural Metropoli- de Cuchorro, 27.VIII.2000). tano, 11.V.1995 (on Pseudobombax septenatum). A single female characterised by strongly microph- Paratenetus grandicornis Motschoulsky, 1868 thalmic head. Probably a new species. (Figs 12–13) Lorelus trapeziderus Champion, 1913 Panama, Panama Prov., Cerro Ancón, 13.I.2002; ídem, 31.I.2002; Parque Natural Metropolitano, 21.III.2001 Panama, Colón Prov., San Lorenzo, Prot. Area, (on (on Luehea seemannii); Colón Prov., San Lorenzo Prot. dead branches of Poulsenia armata in the canopy). Area, 6.III.2001 (on Phryganocydia corymbosa); ídem, 10.III.2001 (on Pourouma bicolor); ídem, 11.III.2001 (on Anaedus corvinus Boheman, 1858 Tovomita longifolia); ídem, 18.III.2001 (on Terminalia amazonica); ídem, 3.V.2001 (on Cydista aequinoctalis); Brazil, Paraná, Foz de Iguassu, 25–30.VIII.2000. ídem, (on Phryganocydia corymbosa); ídem, 21.V.2004 (on Tapirira guianensis); ídem, 31.I.2002 (on Oenocarpus Anaedus maculatus Champion, 1886 mapora); ídem, 4.V.2001 (on Tontelea ovalifolia); ídem (on Forsteronia viridescens); ídem, 10.III.2001 (on Panama, Panama Prov., Gamboa, 10–15.V.1995; Brosimum utile). Ancón, 4.XI.2001, ídem, 5–9.V.2004. 650 J. FERRER and F. 0/DEGAARD

Anaedus marginatus Champion, 1886 Phymatestes agnei sp. nov. (Fig. 1–4, Photo 7) Panama, Panama Prov., Gamboa, 24.V.1995. Brazil. Paraná, Foz de Iguassu, 25–30.VIII.2000. Panama, Panama Prov., Gamboa, 12.VI.1995.

Anaedus setulosus Champion, 1886 Tribe Belopini

Panama, Colón Prov., Colón Prov. San Lorenzo Prot. Adelonia quadricollis (Champion, 1885) Area, 1.VI.2001. Rhacius quadricollis Champion, 1885. Anaedus villosus Champion, 1892 Panama, Colón Prov., San Lorenzo Prot. Area, 8.XI.2001 (on Socratea exorrhiza); ídem, 17.XI.2001 Costa Rica, Guanacaste Prov., Sta Rosa, 11.II.1998; (on Brosimum utile); ídem, 26.IX.2003 (on Poulsenia Panama, Colón Prov., San Lorenzo Prot. Area, 1.VI.2001; armata); ídem, 1.XII.2001 (on Socratea exorrhiza); Panamá Prov., Barro Colorado Isl., 23–31.V.2001. ídem, 28.IX.2003; Panama Prov., Ancón, IX.2001; ídem, 11.IX.2001; Gamboa, 10–15.V.1995. Anaedus apicicornis Champion, 1886 Tribe Nilionini Panamá, Panama Prov., Gamboa, 4–9.V.1995; ídem, 28.V.1995; Summit Graden, 15.IV.1995; Ancón, 1– The knowledge of this group is in poor state. Pic 20.V.1996; ídem, 4.XI.2001; idem, 5–9.V.2004; Brazil, (1918) named new species from Central America and Paraná, Foz de Iguassu, 21–23.VIII.2001. Brazil, giving very short diagnosis. A study of all type material of this author is necessary before description Anaedus sp. of new species.

Panama, Panama Prov., Ancón, 4. XI. 2001 Nilio sp.

Aesymnus nitidus Champion, 1886 Panama, Colón Prov., San Lorenzo Prot. Area. 17.IV.2002 (on Callophyllum longifolium) Panama, Panama Prov., Ancón, 5–9.V.2004; ídem, One of the smallest species from Central America, 4.V.2001; ídem, 2.I.2002. 2.5 mm. long, characterised by strong greenish aeneous colour and by orange-reddish antennae and legs.

Tribe Goniaderini Nilio thomsoni Champion, 1888

Goniadera aff. repanda Fabricius, 1801 Panama, Colón Prov., San Lorenzo Forest, 17.IV.2002. (Fig. 5) (on Callophyllum longifolium); ídem, 28.XII.2001; ídem, 11.X.2003. Goniadera pseudorepanda in litt. = Goniadera repanda Champion, 1886: 236, nec. Fabricius, 1801: 165. Nilio villosus (Fabricius, 1787) Panama, Colón Prov., San Lorenzo Prot. Area, 17. X. 2003. Panama, Colón Prov., San Lorenzo Prot. Area, The examination of the syntypes of Goniadera repan- 24.IX.2003; Panama Prov., Parque Natural Metropoli- da (Fabricus, 1801), described as Melandrya, and all tano, 17.V.1996 (on Anacardium excelsum); available material inclusive types of the Goniadera material preserved in the Natural History Museum, London; Muséum national d´Histoire naturelle, Paris Sufamily Pimeliinae and other Europeans Museums, prove that this new species has been confused with Goniadera repanda. It Tribe Epitragiini will be described in the current revision of the genus Goniadera Perty, in preparation. Bothrotes canaliculatus Say, 1824

Aemymone crenata Champion, 1892 Mexico, San Miguel Negandia, 2200 m a.s.l., 9–22. XII. 1999, Lars Ove Hansen leg.; Panama, Colón Prov., San Lorenzo Prot. Area, 4.V.2001; Nicaragua, 5 Km S of Granada direction towards Panama Prov., Gamboa, 10–15.V.1995; ídem, 24.VI.1995. Nindaime, XII.1999. ídem, Lars Ove Hansen leg.; NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 651

Nicaragua, Granada, 5 Km S of Granada, dirección Tribe Cnemeplatini Nindaline (netted) 1. XII.1999, Lars Ove Hansen leg. The female of this species lack the conspicuous pro- Lepidocnemeplatia laticollis (Champion, 1884) notal carina of the male. Cnemeplatia laticollis Champion, 1884: 136. Cyrtomius plicatus Champion, 1884 Mexico, Estado de Mexico, San Miguel Nepantia, 2200 m a.s.l., 19–22.XII.1999, Lars Ove Hansen leg. Mexico, Estado de Mexico, San Miguel Nepantia, Remarks. According to Kaszab (1938), C. laticollis 2200 m a.s.l., 19–22.XI.1999. Lars Ove Hansen leg. Champion, from Panama and C. sericea Horn, from USA, are synonyms. Phegoneus viridis Champion, 1884

Costa Rica, Guanacaste Prov., Sta Rosa, 11.II.1998. Tribe Asidini

Schoenicus panamensis Champion, 1884 Bothrasida clathrata (Champion, 1884)

Panama, Panama Prov., Parque Natural Metropoli- Asida clathtrata Champion, 1884: 54. tano, 26.II.1996, (on Luehea seemannii); ídem, 18.I.1996 Mexico, Estado de Mexico, San Miguel Nepantia, (on Gouania lupuloides); Ancón, 13.I.2002; Gamboa, 2200 m a.s.l., 19–22.XII.1999, Lars Ove Hansen leg. 23.V.1995.

Epitragus aurulentus Kirsch, 1866 Subfamily Opatrinae

Panama, Panama Prov., Parque Natural Metropoli- Tribe Platynotini tano, 30.IV.1995, (on Cordia alliodora); ídem, 6.IV.1995. Opatrinus gemellatus (Olivier, 1795) Tribe Evaniosomini Panama, Panama Prov., Ancón, 5–9.V.2004. Ditaphronotus foveicollis (Champion, 1884) A widely distributed species in Central America and Antillas. Costa Rica, Guanacaste Prov., Santa Rosa, 11.II.1998; Panama, Colón Prov., San Lorenzo, 3.I.2002; Nicaragua, Managua, Carretera Masaya, 12.5 km, netted, XI– Tribe Opatrini XII.1999, Lars Ove Hansen leg. The numerous species of Blapstinus Sturm, 1826, Hylocrinus parallelus (Champion, 1884). lack an accurate revision, initiated by Davis (1970) and still in course. We include some undeterminable species, Emmenastus parallelus Champion 1884: 12 to attire the attention of specialists.

México, Estado de Mexico, San Miguel Nepantia, Blapstinus metallicus (Fabricius, 1801) 19–22.XII.1999, 2200 m a.s.l., Lars Ove Hansen leg. Panama, Panama Prov., Gamboa, 30.XI.1995; Panama Tribe Branchini Prov., Ancón, 11.XI.2001.(3); ídem, 4.XI.2001; ídem.10. III.1995 (2). Branchus obscurus Horn, 1866 Blapstinus grandis Champion, 1885 Nicaragua, 5 km S of Granada, direction to Nindaime, 1.XII1999 (netted) Lars Ove Hansen leg. Nicaragua, Granada, 2 km of Nindaime (netted at dry river bed) 1.XII.1999, Lars Ove Hansen leg.; Costa Tribe Stenosini Rica, Sta Elena, 10.II.1995.

Rhypasma livae sp. nov. Blapstinus buqueti Champion, 1885 (Figs 38–40, Photo 8) Panama, Panama Prov., Gamboa, 3.XII.1995; ídem, Panama, Panama Prov., Ancón, 16.II.2002, A. 4.XI.2001; ídem, 4–9.V.1995; ídem, 9.I.1996, Panama Ødegaard leg. Prov., Ancón, 4.XI.2001; ídem, 11.XI.2001. 652 J. FERRER and F. 0/DEGAARD

Blapstinus errabundus Champion, 1885 A single male, Panama, Panama Prov., Parque Natural Metropolitano, 7.V.1996 (on Enterolobium cyclocarpum); Panama Prov., Gamboa, 10–15,V.1995; ídem, a female: Panama, Panama Prov., Gamboa, 5.VI.1995. 26.V.1995; Panama Prov., Ancón, 4.XI.2001 (3); the Gebien (1939) and Papp (1961) considered C. aspe- specimens are somewhat different and represent per- rulus Pascoe as a synonyme of cucullatus. However, haps two local populations of same species. Champion (1886) considered asperulus as a form or variety of cucullatus Pascoe, characterised by apically Blapstinus sp. 1 divergent precephalic horn, which is shortly truncate in the nominal form (Figs 50–51). In fact, C. asperulus Costa Rica, Guanacaste Prov., Sta Rosa, 11.II.1998. is the male of C. cucullatus, which described by Pascoe after a female in an anterior page, is prioritary. Blapstinus sp. 2 Tribe Diaperini Nicaragua, Granada, 2 km of Nindaime (netted at dry river bed) 1.XII.1999, Lars Ove Hansen leg. Liodema serricorne Bates, 1873

Blapstinus sp. 3 = Liodema flavovariegatum Champion, 1886: 208. Panama, Colón Prov., San Lorenzo Prot. Area, Panama, Panama Prov., Parque Natural Metropoli- 19.I.2002 (on Brosimum utile). tano, 7.V.1996 (on Serjania mexicana). Platydema guatemalensis Champion, 1886 Blapstinus sp. 4 Panama, Colón Prov., San Lorenzo Prot. Area, Brazil, Paraná, Foz de Iguassu, 25–30.VII.2000 (3). 27.V.2004, Cornejo et al. leg.

Ulus hirsutus Champion, 1885 Platydema undata (Chevrolat, 1878)

Costa Rica, Guanacaste Prov., Sta Elena, 10.II.1999. Panama, Panama Prov., Gamboa, 25.V.1995; ídem, 10–15.V.1995. Ulus lineatulus Champion, 1885 Platydema quinquedecimmaculatum Chevrolat, 1878 Panama, Panama Prov., Gamboa, 20.I.1995; Cerro Ancón, 13.I.2002, F. and A. Ødegaard leg. Panama, Panama Prov., Barro Colorado Isl. 20.IX– 30.X.2003 (2). Trichoton curvipes Champion, 1885 Platydema tibiale Chevrolat, 1878 Panama, Panama Prov., Ancón, 10.III.1995; ídem, 4.II.2002, ídem, 5–9.V.2004. Panama, Panama Prov., Barro Colorado Isl. 20.IX- –30.X.2003. Subfamily Diaperinae Platydema panamensis Champion, 1886 Tribe Dysantini Panama, Panama Prov., Barro Colorado Isl. 20.IX- Ozolais elongata Champion, 1886 –30.X.2003. (Fig. 54) Platydema unicolor Champion, 1886 Panama, Panama Prov., Parque Natural Metropoli- tano, 21.XII.2001; ídem, 14.V.1996; Barro Colorado Isl. Panama, Colón Prov., San Lorenzo Prot. Area, 11–13.V.2004. 1.II.2002 (on Brosimum utile). Remarks. According Doyen and Tschinkel (1982) this genus belongs to Toxicini. Platydema sp. aff. sobrinum Chevrolat, 1877 (Fig. 47) Calymmus cucullatus Pascoe, 1871 (Figs 50–53) Panama, Colón Prov., San Lorenzo Prot. Area, 20.IX– 30.X.2003 (2). = Calymmus asperulus Pascoe, 1871: 350. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 653

The numerous species of Platydema north of Mexico Antimachus coriaceus Lacordaire, 1859 has been revised by Triplehorn (1965). Philips et al. (Figs 55–59) (1998) described P. woldai, a new species from Panama, Guatemala and Honduras. The Central American rela- Panama, Panama Prov., Parque Natural Metropoli- tives are still waiting for a review after the classical work of tano, 6.V.1995 (male); ídem, 4.IX.1995. Champion (1886). This species is closely related to P. nigra- This species recorded by Champion (1886), from tum Motschoulsky, 1873 and P. sobrinum Chevrolat, 1877, Nicaragua to Brazil, has been overlooked by Gebien but the aedeagus is different. It is impossible to key using (1940) in his Catalogue. Champion (1886) or Triplehorn (loc. cit.). The species is highly probably undescribed, however creating a new Peneta nuchicornis Gebien, 1928 name to add to the list of 91 described Central American taxa of Platydema, before an accurate examination of Panama. Colón Prov., San Lorenzo Prot. Area, 20. IX available type material of at least all species exhibiting dark – 30. X. 2003, A. Tishechkin leg. opaque body and more or less reddish legs and antennae, Remarks. Peneta nuchicornis Gebien (1928) has been seems to be premature. We give a short diagnosis and fig- described from Costa Rica and P. nuchicornoides Kaszab ures of this species: dimensions: 7.4 mm × 4.5 mm; body 1977, from Colombia, as figured in Kaszab (1977). New ovate, dull, pruinous opaque, with reddish antennae and record for Panama. legs, dorsally practically unpunctured, except the head, which is unarmed in both sexes; the tegument shiny and Sitophagus dilatifrons Champion, 1886 densely covered of nearly contiguous, round punctures. Ventral surface blackish, shiny and superficially punc- Panama, Panama Prov., Parque Natural Metropoli- tured. Aedeagus small and pointed apically (Fig. 47). tano, 12.II.1995 (male).

Paniasis kulzeri sp. nov. Hypogena tricornis (Laporte de Castelnau, 1840) (Figs 41–46, Photo 6) Costa Rica, Guanacaste Prov., 11.II.1998 (male). Panama, Panama Prov., Gamboa, 10.V.1995. Remark. The old genus Ulosonia (Laporte de Castelnau 1840: 220), recently revised by Kulzer (1962) Cosmonota silphoides (Laporte de Castelnau) comb. nov. has been changed to Hypogena.

Platydema silphoides Laporte de Castelnau, 1831: 369. Alegoria dilatata (Laporte de Castelnau, 1840)

Panama, Colón Prov., San Lorenzo Prot. Area, Panama, Panama Prov., Gamboa, 4–8.V.1995; Parque 21.XII.2001 (on Cassipurea elliptica). Natural Metropolitano, 1.V.1995, Panama Prov., Ancón, 5–9. This insect, abnormally large and depressed for the XI.2001; Colón Prov, San Lorenzo Prot. Area, 18.V.2001. genus, has been described as Platydema, but exhibits the peculiar V-shaped metasternum and the shape of Iccius cephalotes Champion, 1886 Cosmonota, claiming a transference to this genus. ♂, Panama, Colón Prov., San Lorenzo Prot. Area, Tribe Phrenapatini 22.IX–30.IX.2000, IBISCA team leg.; ♂, ídem, 6.III.2001 (on Brosimum utile); ♀, ídem, Phrenapates bennetti Kirby, 1837 24.V.2001; ♂, ♀, ídem, 19–X.2001; ♀, ídem, 22.IX.2001 (on Poulsonia armata); ♂, 24.XI.2001 (on Brosimum Panama, Chiriqui Prov., 2 km north of Fortuna, 1100 utile); ♂, ídem, 22.XII.2001 (on Brosimum utile); ♀. m a.s.l., 19.VI.1995, A.R. Gillogly leg. Recorded from ídem, 24.I.2002 (on Brosimum utile); ♀, Mexico, Estado Mexico to Colombia. de Mexico, San Miguel Nepantia, 2200 m a.s.l., 19- The aedeagus of the specimen from Panama is prac- 22.XII.1999, Lars Ove Hansen leg. tically identical to the nominal form, from Colombia. However, the cephalic horn is much shorter and massive. Iccius monoceros sp. nov. (Figs 48–49) Tribe Ulomini ♂, Panama, Colón Prov., San Lorenzo Prot. Area, Uloma retusa var. dimidiata Champion, 1886 13.X.2003, R. Kitching leg. (at lights); ídem, Flight intercepted trap, canopy, 21 m above forest floor, 25.V–4.VI.2004, R. Panama, Panama Prov., Parque Natural Metropoli- Didham et al.; ídem, 1–13.X.2003, R. Didham and L. Fagan tano, 24.VIII.1995. leg. Museum of Natural History, Budapest, ♂. ♀. 654 J. FERRER and F. 0/DEGAARD

Tribe Triboliini Nuptis corticalis Champion, 1885

Tribolium confusum Jacquelin du Val, 1868 Panama, Chiriqui Prov., Boquete, 1200 m a.s.l., 8. VIII. 1995. Panama, Panama Prov., Ancón, 5.IV.1995; ídem, 11.IX.2001; ídem, X.2001. Nuptis tenebrosa Champion, 1885 Remark. A cosmopolite, widely distributed pest of stored products. Panama, Colón Prov., San Lorenzo Prot. Area, 16.XI.2001 (on Brosimum utile).

Subfamily Tenebrioninae Hesiodus cf. conspurcatus Champion, 1885

Tribe Alphitobini Panama, Panama Prov., Gamboa, 21.V.1995; Colón Prov., San Lorenzo Prot. Area, 29.XII.2001 (on Symphonia Alphitobius laevigatus (Fabricius, 1781) globulifera); ídem, 25.IV.2002; ídem, 11.V.2004 (on Pouroma bicolor). Helops laevigatus Fabricius, 1781: 90. This species is similar, but not exactly corresponding Panama, Panama Prov., Ancón, 10.IV.1995. to the characters, invocated by Champion (loc. cit.), to A cosmopolite species. distinguish the five known species of Hesiodus inhabit- Remark. According Doyen and Tschinkel (1982) ing Central America. The body is black, pruinose, the Alphitobius belongs to Tenebrionini. striae of elytra strongly punctured, the intervals very finely and sparsely punctured and the humeri dentiform. The single male specimen examined is insufficient for a Tribe Tenebrionini decision concerning the specific status of this insect.

According Doyen and Tschinkel (1982) the sub- Hesiodus cf. longitarsis Champion, 1885 family Tenebrioninae has to include the new tribe Coelometopini. The position of the genus Nuptis Panama, Panama Prov., Barro Colorado Isl. 11–13. Motschoulsky 1872 and other genera cited here remain V. 2004. uncertain into comparative study. As in the precedent case, the identification of this species remains uncertain. It is impossible to assign this Zophobas opacus (Sahlberg, 1923) single female a specific or subspecific status.

Tenebrio opacus Sahlberg, 1823: 17. Choastes purpureus (Champion, 1885) Panama, Panama Prov., Gamboa, 6.I.1995. Choaspes purpureus Champion, 1885: 119 Zophobas tridentatus Kraatz, 1880 Panama, Colón Prov., San Lorenzo Prot. Area, 10.XII.2001. Panama, Panama Prov., Parque Natural Metropolita- no, 22.V.1995 (male); Panama Prov., Barro Colorado Isl., Choastes angulicollis (Champion, 1885) 20:IX–30.X.2003 (female). Remark. The taxonomy and synonymy of the genus Choaspes angulicollis Champion: 1885: 119 Zophobas Blanchard, is one of the almost intricate. The Panama, Panama, Colón Prov., San Lorenzo Prot. recent discovery (Ferrer et al. 2004) of lost type material Area; ídem, 8.V.2001; ídem, 27.V.2001; ídem, 28.IX.2001; from Fabricius and Olivier, preserved in the William ídem, 14.X.2001; ídem, 23.XI.2001 (on Brosimum utile); Hunter collection, of the Hunterian Museum of the ídem, 25.IV.2002 (on Brosimum utile) University of Glasgow, facilitate the identification of This genus is unambiguous by the dentate underside these species, which has been compared with all types. of posterior femora. Kulzer (1964) has described two species from Peru. Mylaris procera (Champion, 1885)

Nyctobates procera Champion, 1885: 107. Tribe Coelometopini Panama, Panama Prov., Parque Natural Metropoli- tano, 7.V.1995. [Doyen and Tschinkel (1982), pars. = Cnodalonini The genus Nyctobates Guérin, 1834: 33, has been Gebien] found identical to Mylaris Pallas, 1781. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 655

Othryoneus triplehorni sp. nov. Brosimapsida gonospoides gen. et sp. nov. (Figs 74–76, Photo 1) (Figs 69–73, Photo 14)

Panama, Colón Prov., San Lorenzo Prot. Area, Panama, Colón Prov., San Lorenzo Prot. Area, 11. V. 2004. 21.XII.2001 (on Brosimum utile); ídem, flight-intercept trap, 35 meters above ground, 1–17.X.2003, R. Lenkous ibisca sp. nov. Didham and L. Fagan leg. (Museum of Natural History, (Figs 33–37, Photo 16) Budapest).

Holotype: ♂, Panama, Colón Prov., San Lorenzo Cyrtosoma denticollis Chevrolat, 1878 Prot. Area, extracted by Berlese-Tullgren from epiphytes collected in the canopy, 27.IX.2003, N. Winchester and Panama, Panama Prov., Gamboa, 3.X.1995; Colón K. Jordan leg. Paratypes: (sex not examined) Panama, Prov., San Lorenzo, 28.XII.2001; ídem, 6.V.2002, Colón Prov. San Lorenzo Prot. Area, 8.X.2004, J. Bail leg. A. Tisechkin leg. ; 5.IX.2002; ídem, 20.IX–30.X.2003. (canopy fogging) Museum of Natural History, Budapest and coll. J. Ferrer. Cyrtososma decimlineata Champion, 1886

Gonospa phaedonoides Champion, 1886 Panama. Panama Prov., Ancón, 5.IV.1995. (Figs 66–68) Cyrtosoma sp. cf. decemlineata Champion, 1886 Panama, Panama Prov., Parque Natural Metropolitano, 29.I.1996; ídem, 6.V.2004; Colón Prov., San Lorenzo Panama, Panama Prov., Barro Colorado Isl., 2. V. 2004. Prot. Area, 21.X.2001; ídem, 3.IV.1995; ídem, 6.IV.1995. This species is similar to decemlineatum, but the disposition of entirely coloured intervals is different. Gonospa similis sp. nov. In decemlineatum the suture and the alternate intervals (Figs 60–62, Photo 13) are red to orange. In this single specimen the suture, the 3.4.5.7 and the lateral margin are black, however the Panama, Panama Prov., Parque Natural Metropolitano, black bands are interrupted basally and apically and the 24.IV.1995 (on Bonamia marapioides); ♀, ídem: patch of the 5 interval is very reduced. Using Marcuzzi’s 2.II.1995 (on Dioclea guianensis); ídem, 15.I.1996; ídem: key (1991) this specimen has to be placed between lin- 5–6.IV.2004; San Lorenzo Forest, IBISCA, Teneb65, eatum Laporte de Castelnau, 1831, from the Amazonas Museum of Natural History, Budapest. region and decemlineatum. However, the coloured bands of lineatum are limited to the lateral intervals, sometimes Apsida simulatrix sp. nov. reduced: the South American var. reductevittatum Pic, (Figs 63–65, Photo 15) 1935 of lineatum. A new revision, based in genital characters and comprehensive figures of this specious and Panama, Colón Prov, San Lorenzo Prot., Area, unfortunately poorly known genus, is necessary. 4.V.2001 (on Virola elongata); ídem, 3.IX.2003, H. Barrios and F. Ødegaard leg.; ídem, 11.V.2002 (on Cyrtosoma sp. Arrabidaea verrucosa); ídem, 30.XI.2001, (on Marila laxifolia); ídem, 24.I.2002. Panama, Panama Prov., Gamboa, 4–9.V.1995, Colón Prov., San Lorenzo Prot. Area, 26.IV.2002 (on Dussia Apsida purpureomicans Bates, 1873 munda). Another red banded species, exhibiting strongly Panama, Panama Prov., Parque Natural Metropoli- carinate protibia. Probably a new species. As both tano, 5.II.1995 (on Enterolobium cyclocarpum); ídem, specimens are females it seems premature to create new 21.VIII.1995 (on Enterolobium cyclocarpum); ídem, names before revision of the genus Cyrtsosoma. 27.IX.1995; ídem, 5–6.V.2001; Panama Prov., Cerro Ancón, 13.I.2002. Epicalla sp. nov. (Fig. 79) Apsida boucardi Bates, 1873 Ecuador, Santo Domingo de Colorados, G. Onoré Panama Prov., Parque Natural Metropolitano,10.VI. (male). Panama, Colón Prov., San Lorenzo Prot. Area, 1995, (on Mikania leiostachya); ídem, 22.V.1995; Cerro 23. XI. 2001. Ancón, 13.I.2002. The species of the genus Epicalla Champion, 1886, are recognised by 7–8 striae of elytra forming a strong 656 J. FERRER and F. 0/DEGAARD premarginal carina. This widely distributed species has to tened discally, finely sculpted of superficial and fine rows be described in a separate paper revising the Amazonian of punctures becoming carinate at sides and posteriorly, representatives of the genus Epicalla Gebien 1938. the intervals separated by clearly traceable carinate elevations, between the rows of fine punctures. Epicalla aeneipes sp. nov. Ventral zone, glabrous, except the coxae, shining (Fig. 84, Photo 10) green metallic, mentum strongly trapezoidal, shiny, convex, with some scarce hairs, gula convex, deeply separat- Panama, Panama Prov., Parque Natural Metropoli- ed by a transversal impression; propleura shiny, polished tano, 17.IV.1995; San Lorenzo Forest, IBISCA, 12.V.2004, and concave; prosternum strongly punctured, proster- Museum of Natural History, Budapest). nal apophysis triangular, flattened an finely impressed at middle. Mesosternum carinate at middle and rugose Epicalla elongata sp. nov. at sides. Metasternum broadly opened in “V”, to receive (Figs 77–78, 86, Photo 9) the apophysis, concave at middle, forming a strongly depressed zone, epimeral, episternal and epipleural Panama, Colón Prov., San Lorenzo Prot. Area, zones deeply, irregular and strongly punctured. 11. V. 2001 (on Dendropanax arboreus). Epicalla cupreonitens Champion, 1886 Epicalla pygmaea sp. nov. (Fig. 85, Photo 12) Panama, Panama Prov., Parque Natural Metropoli- tano, 17.IV.1995, (2) (on Cordia alliodora); ídem, Panama, Panama Prov., Parque Natural Metropoli- 6.V.1995 (2) (on Antirrhoea trichantha). tano, 4. VI. 1995. Camaria parallela Champion, 1886 Epicalla instriata Pic, 1921: 28 (Figs 81–83, 87, Photo 11) Panama, Panama Prov., Parque Natural Metropoli- tano, 19.X.1995 (on Arrabidaea corallina); ídem, Panamá, Colón Prov., San Lorenzo Prot. Area, 29.I.1996 (on Astronium graveolens); Colón Prov., San 15. XII. 2001. Lorenzo Prot. Area, 9.III.2001; ídem, 1.IX.2001, (on This species has been overlooked in the Gebien Socratea exorrhiza); ídem, 5.XII.2001. (1928) revision. The laconic diagnosis of Pic (1921) has to be completed as following. Calydonella lisa Doyen, 1995 Length: 14 mm, maximum of width: 6 mm. (Fig. 80) Head broadly rounded anteriorly, macrophthalmic; epistome finely impressed, labrum subquadrate, punc- Panama, Chiriqui Prov., Fortuna, 8.VIII.1995. A sin- tured and pubescent, supra-antennal zones strongly fove- gle male from the locus tipicus. ate before eyes, which are strongly globose and separated Remark. Doyen (1995) never studied the genitalia frontally by a distance equal to two time the diameter of this monotypic genus. We have examined for first of an eye measured dorsally, becoming equal posteri- time the aedeagus of this insect (Fig. 80). He placed orly. The eyes are reniform in lateral view, as results of Calydonella near Othryoneus Champion, 1886. However, the inserction of the ocular canthus; tegument finely, examination of type material of all species of this genus sparsely and superficially punctured. Antennae reaching revised by Kulzer 1964), indicate that Othryoneus are the shoulders, third joint a little longer than the 4th, the vively coloured recalling the decoration of Diaperis following becoming larger, but decreasing to the apex and boleti (L.) and other Diaperini, Calydonella is a metallic becoming lighter apically as indicated in Fig. 83. insect and in all essential characters agree and has to be Pronotum feebly transverse, broadly rounded ante- placed near the genus Calydonis Pascoe, 1882. riorly, without conspicuous anterior angles, the sides becoming subsinuated and parallel before the base, pos- Blapida alternata Gebien, 1919 terior angles right, strongly carinate, with a little fovea at the middle of the anterior margin and the middle Panama, Colón Prov., San Lorenzo Prot. Area, of the base, which exhibit two depressed fovea at each 22.XII.2001 (on Bromium utile); ídem, 9.III.2001; ídem, side; tegument finely and sparsely punctured. Scutellum 15.XII.2001. triangular, metallic green, shiny, polished. Elytra strongly convex, subglobose in profile, 1.5 times Acropteron belti Champion, 1886 as long as broad, the shoulders strongly callose and carinate, the humeral carina surpassing the level of the base, Panama, Panama Prov., Barro Colorado Isl. 20.I.2004; which is profoundly depressed at each side; tegument flat- Parque Natural Metropolitano, 21.XII.2001. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 657

This remarkable species is characterised by a post- Nautes sp. 4 genicular plaque covering the first third of the metatibia of the female. The metatibia of the male are subsinuate. Panama, Panama Prov., Gamboa, 3.X.1995; Parque Natural Metropolitano, 4.X.1995 (on Arrabidaea patel- Acropteron languroides Champion, 1886 lifera).

Panama, Panama Prov., Parque Natural Metropoli- Tribe Talanini tano, 21.XII.2001; Colón Prov., San Lorenzo Prot. Area, 9.V.2002 (on Cassipourea elliptica). Talanus neotropicalis Champion, 1887 This species is hardly separable from the A. belti. Perhaps only a much finer punctured subspecies. Panama, Panama Prov., Pipeline Road, 25.I.2002; Parque Natural Metropolitano, 8.VII.1995 (on Gouania Acropteron longipenne Champion, 1886 lupuloides); ídem, 6.XI.1995 (on Serjania rhombea).

Panama, Panama Prov., Cerro Campana, 650 m a.s.l., Tribe Strongyliini 16.V.1996 (2); Panama, Colón Prov., San Lorenzo Prot. Area, 15.V.2001. Strongylium vikenae sp. nov. A. longipenne is the only species exhibiting simple (Figs 88–90, Photo 2) metatibia. Panama, Panama Prov., Barro Colorado Isl. 11– Xenius scabripennis Champion, 1886 13.V.2004 and 22–31.V.2004; San Lorenzo Prot Area, 9–28.V.2004, G. Curletti leg. (2) Museo Civico de Panama, Panama Prov., Parque Natural Metropoli- Carmagnola and coll. J. Ferrer). tano, 2.XII.1995; Panama Prov., Parque Natural Soberanía, Plantation Road, 9.XII.2001. Strongylium angustulum Mäklin, 1864

Tribe Helopini Panama, Panama Prov., Cerro Campana, 650 m a.s.l., 16.V.1995. Nautes breviceps Champion, 1886 Strongylium aulicum Mäklin, 1864 Panama, Colón Prov., San Lorenzo Prot. Area, 22.XII.2001 (on Brosimum utile); ídem, 21.I.2002, (on Panama, Colón Prov., San Lorenzo Prot. Area, Brosimum utile); ídem, 24.2002 (from dead branches of 22.VIII.2002; ídem, 8.V.2001; ídem, 18.V.2001; ídem, Brosimum utile in the canopy). 24.V.2004; Panama Prov., Gamboa, 28.V.1995. Remarks. Five species unfortunately only represent- ed by female specimens. This specious, poorly known Strongylium aff. blandum Mäklin, 1864 genus needs an accurate revision before description of new taxa. Panama, Colón Prov., San Lorenzo Prot. Area, 18.V.2001 (on Vantanea depelta). Nautes sp. 1 Strongylium brevipes Champion, 1888 Panama, Colón Prov., San Lorenzo Prot. Area, 22: XII.2001 (on Brosimum utile), idem, 22.I.2002 (on Panama, Colón Prov., San Lorenzo Prot. Area, Clusia aff. longipetiolata). 14.V.2002 (on Dendropanax arboreus); ídem, 8.V.2001; ídem, 11.V.2001; ídem, 24.V.2004; Panama Prov., Parque Nautes sp. 2 Natural Metropolitano, 24.IV.1995; ídem, 26.VI.1995.

Panama, Panama Prov., Parque Natural Metropo- Strongylium decoratum Mäklin, 1864 litano, 11.XII.1995 (on Pithecoctenium crucigerum); Barro Colorado Isl. 20.IX.–30.X.2003. Panama, Panama Prov., Parque Natural Metropoli- tano, 26.VI.1995 (on Cordia alliodora); Nautes sp. 3 Colón Prov., San Lorenzo Prot. Area, 29.V.2004, idem, 4.V.2001; ídem, 8.V.2001; ídem, 11.V.2001; ídem, Panama, Colón Prov., San Lorenzo Prot. Area, 25.V.2001; ídem, 25.IV.2002; 31.I.2002 (on Inga cocleensis). From Costa Rica to Bolivia. 658 J. FERRER and F. 0/DEGAARD

Strongylium delauneyi Flétiaux et S., 1889 Strongylium sp. 2 (Photo 18) Panama, Colón Prov., San Lorenzo Prot. Area, 20– 27.V.2004, Cornejo et al. leg. Pamamá, Colón Prov., San Lorenzo Prot. Area, A rare species described from Guadaloupe. New 7.V.2004; Panama Prov., Parque Natural Metropolitano, record for Panama. 8.V.1995 (on Cordia alliodora), Barro Colorado Isl., 11–13.V.2004. Strongylium exaratum Champion, 1887 A macrophthalmic, green metallic, aeneous species, with alternate reddish brown tint on the shining inter- Panama, Colón Prov., San Lorenzo Prot. Area, vals, only visible examining the insect from lateral view. 31.V.2004 (on Apeiba membranacea); ídem, 15.V.2004. 10.5 mm long, with slender antennae and legs.

Strongylium excavatum Mäklin, 1864 Strongylium sp. 3 (Photo 19) Panama, Panama Prov., Gamboa, 10.V.1995; Barro Colorado Isl. 11–15.V.2004.; Panama Prov., Parque Panama, Panama Prov., Gamboa, 11.VI.1995. Natural Metropolitano, 8.V.1995, ídem, 9.V.2004. A cupreous-aeneous species with green metallic tint on the epistoma, the legs and antennae brownish black, Strongylium ignitum Champion, 1887 the body relatively broad and short, 9.3 mm long.

Panama, Colón Prov., San Lorenzo Prot. Area, Strongylium sp. 4 4.V.2001 (on Symphonia globulifera); ídem, 4.V.2001 (on (Photo 20) Apeiba membranacea); ídem, 8.V.2001; ídem, 3.V.2002 (on Brosimum utile); ídem, 15.V.2004; ídem, 18.V.2001 Panama, Panama Prov., Parque Natural Metropoli- (on Guatteria dumetorum). tano, 26.IV.1996 (on Maeluta tinctoria). Previously only recorded from Nicaragua. Another short and massive species, aeneous, with short antennae and legs. 7.3 mm long, well characterised Strongylium sp. aff. interstitialis Germar by contiguous, superficially, but finely incised foveate, round circles, replacing punctures on pronotum. Panama, Panama Prov., Gamboa, 30.XI.1995, Colón Remarks. Many small species of the genus Strongy- Prov., San Lorenzo Prot Area, 29.V.2004. lium sensu Mäklin 1884 can not be satisfactory studied, before an accurate examination of all available Strongylium aff. leprosum Mäklin types of Mäklin, unfortunately dispersed in Stockholm, Helsinki, Vienna and Berlin Museums. As in the case Panama, Colón Prov., San Lorenzo Prot. Area, of Blapstinus, we give a number and a short diagnose 4.V.2002 (on Dendropanax arboreus); ídem, 2.VI.2004 of four unnamed, highly probably new species of small (on Terminalia amazonica); ídem, 8.V.2001 (on Pera Strongylium in order to attire the attention of future arborea). specialists.

Strongylium panamense Champion, 1888 Otocerus angelicae sp. nov. (Figs 93–95, 97–98, Photo 4) Panama, Panama Prov., Gamboa, 27.V.1995; Colón Prov., San Lorenzo Prot. Area, 25.V.2001, ídem, 23.V.2004 Panama, Panama Prov., Barro Colorado Isl., 11– (on dead branches in the canopy). 13.V.2004.

Strongylium sp.1 Otocerus delicatus sp. nov. (Photo 17) (Figs 91–92, 99–100, Photo 3)

Panama, Panama Prov., Parque Natural Metropoli- Panama, Panama Prov., Parque Natural Metropoli- tano, 6.V.1996 (on Ficus maxima); ídem, 10.V.1996 (on tano, 5–6.V.2004. Ficus insipida). A purplish species, with orange femora and distal Cuphotes elongatus (Thomson, 1859) tibial zone, the rest and the tarsi black with a metallic tint. The antennae black, the ventral surface blackish Panama, Panama Prov., Parque Natural Metropoli- brown. tano, 25.IV.1995. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 659

Poecilesthus nigropunctatus Champion, 1887 Poecilesthus sensu lat. It is premature to name this probably new species, before a revision of both extremely Panama, Colón Prov., San Lorenzo Prot. Area, specious genera. 9.III.1995; Panama Prov., Barro Colorado Isl., 20.V.2001. Subfamily Monommatinae

Poecilesthus fragilicornis Champion, 1887 The definitive position of this group is under study. The current place between Zopheridae is questionable. Panama, Colón Prov., San Lorenzo Prot. Area, 15.V.2001 (on Dendropanax arboreus); ídem, 24.V.2004; Hyporrhagus emarginatus Champion, 1888 ídem, 11.V.2002. Panama, Colón Prov., San Lorenzo Prot. Area, Poecilesthus latus Champion, 1887 15.V.2001 (on Dendropanax arboreus); ídem, 29.X.2001, (on ídem). Panama, Colón Prov., San Lorenzo Prot. Area, 11.V.2002 (on Pourouma bicolor); ídem, 20.IV.2002. Hyporrhagus suturalis Champion, 1888

Poecilesthus immaculatus Champion, 1887 Panama, Colón Prov., San Lorenzo Prot. Area, 30.XI.2001 (on Carapa guianensis); Costa Rica, Estación Panama, Colón Province, San Lorenzo Prot. Area, de Biologia, La Selva, VIII–IX.1994, K. Thunes leg. 29.V.2004. Hyporrhagus ferrugineus Champion, 1888 Poecilesthus cupripennis Champion, 1892 Panama, Gamboa, 4–9.V.1995; Colón Prov., San Panama, Colón Prov., San Lorenzo Prot. Area, Lorenzo Prot. Area, 20.II.2001 (on Virola elongata). 11.V.2001 (on Brosimum utile); ídem, 18.V.2001 (on Guatteria dumerorum); ídem, 21.V.2004 (on Terminalia Hyporrhagus nitidus Champion, 1888 amazonica). Panama, Colón Prov., San Lorenzo Prot. Area, 4.V.2001 Poecilesthus variipes Champion, 1887 (on Apeiba membranacea); Panama, Prov., Parque Natural (Figs 101–102) Metropolitano, 11.V.1995 (on Enterolobium cyclocarpum).

Panama, Panama Prov., Gamboa, 11.VI.1998. Hyporrhagus sp. The characters of this species seem to be a combination of Poecilesthus and Strongylium. The aedeagus Panama, Panama Prov., Gamboa, 7.VI.1995; Parque (Figs 101–102) of P. variipes is quite different from Natural Metropolitano, 8.V.1995 (on Astronium gra- other African and Neotropical Strongylium and sup- veolens). port a generic separation from Poecilesthus and from Strongylium when both speciose genera will be studied. Aspathines ovatus Champion, 1888

Genus incertae sedis Panama Prov., Parque Natural Metropolitano, 31.VI.1995 (on Arrabidaea patillifera); Brazil: Paraná, Strongylium? sp. clavicornis group Foz de Iguassu, 2–IX.2000. (cf Figs 103 and 104, Photo 21)

Two females: Panama, Colón Prov., San Lorenzo Discussion Prot. Area, 2.VI.2001. Parque Natural Metropolitano, 13.V.1995 (on Bignoniaceae sp.). As this is one of the first studies of a broad col- A remarkable, highly probably new species, undoubt- lection of Tenebrionidae in tropical forests of Central edly related to Strongylium clavicornis Champion, 1892, America for about hundred years, it is expected that which according to the original description and figure several undescribed species of the beetles appear even in is different from all the allied forms by the short and samples achieved with commonly used collecting meth- strongly clavate antennae. S. clavipes is greenish, metal- ods. However, the investigation of canopy habitats is obvi- lic with black, feebly violaceous elytra. This species is ously an additional important source for discovery of new smaller, 5 mm. long, shorter, shining castaneous brown species of Tenebrionidae in the present study. Species like with reddish appendages. Habitually it seems closer to Lenkous ibisca, Iccius monoceros, Gonospa similis, Apsida 660 J. FERRER and F. 0/DEGAARD simulatrix, Brosimapsida gonopoides, Otocerus delicatus Blackwelder, R. E. 1945. Checklist of Coleopterous Insects of Mexico, and O. angelicae, the three new Epicalla species, as well Central America, the West Indies and South America, Part 3, USA, National Museum, Bulletin, 185: 343–550. and many Strongylium and Poecilesthus species are all most Champion, G. C. 1884–1888. Biologia Centrali-Americana. Insecta, likely restricted to habitats more frequently occurring in Coleoptera (Heteromera). Godman and Salvin eds., London. the canopy than on ground. A significant component of Vol. 1, 476 pp canopy dwellers is constituted by Tenebrionidae although Champion, G. C. 1892. Supplement of Biologia Centrali Americana, not as prominent as in phytophagous taxa. For this reason IV. Insecta. Coleoptera. (Heteromera) Godman and Salvin eds., London. 477–572. 23 pl. also future studies should be focused on canopy habitats. Champion, G. C. 1913. Notes on various Central American Coleoptera Regarding host relationships, the Tenebrionidae show with descriptions of new genera and species. Transactions of the low degree of preference for certain host plants. This Entomological Society of London, 58–169. is expected for detritivorous and fungivorous species, Champion, G. C. 1917. On new and little known Lagriinae from although some species may be strongly associated with tropical America. Coleoptera. Transactions of the Entomological particular species of fungi. The fungi themselves, howev- Society of London, 169–267. Davis, J. C. 1970. Revision of the genus Blapstinus Sturm of America er, tend to be generalists in the tropics (Ødegaard 2000b). North of Mexico with notes on extralimital species (Coleoptera, Based on large samples just a few beetle species show Tenebrionidae). Thesis. The Ohio state University, 459 pp. preferences to host plants in this study: Iccius cephalotes (unpublished). on Moraceae and Blapida alternata on Brosimum utile. Doyen, J. T. 1988. Tenebrionidae and Zopheridae of the Chamela Biological Station and vicinity, Jalisco, Mexico (Coleoptera). Folia Entomologica Mexicana, 77: 211–276. Doyen, J. T. 1989. Reconstitution of the Diaperini North of Mexico, Acknowledgements with new species of Adelina and Sitophagus. Coleoptera, Tenebrionidae. Proceedings of the Entomological Society of We are grateful to the canopy Crane Team at Washington, 86 (4): 777–789. STRI, including S. J. Wright, Y. Basset, V. Horlyck, M. Doyen, J. T. 1995. A new genus and four new species of Coelometopini from Mesoamerica (Coleoptera, Tenebrionidae). The Coleopte- Ruiz, M. Samaniego, J. Herrera, E. Andrade and O. rist´s Bulletin. 49 (1): 8–14. Saldana. Great thanks to Åslaug Viken, for field assist- Doyen, J. T., Matthews, E. and J. F. Lawrence. 1989. Classification and ance and to Oddvar Hanssen, NINA for help with Annotated Check-list of the Australian genera of Tenebriondae. photos. Unvaluable help with type consultation has Coleoptera, Invertebrate taxonomy, 3: 229–260. been provided by M. Max V. L. Barclay, The Natural Doyen, J. T. and W. Tschinkel. 1982. Phenetic ad Cladistic relationship about Tenebrionid beetles. Coleoptera, Systematic Entomology, History Museum (SYSRESOUR), London, Dr. Claude 7: 157–183. Girard, Muséum national d´Histoire naturelle, Paris Erwin, T. L. 1982. Tropical forests, their richness in Coleoptera and (COLPARSYS), Dr. Dariusz Iwan, Museum and Institute other arthropod species. The Coleopterist’s Bulletin, 36: 74–75. of Zoology, MIZ, Warszawa (SYNTHESYS), Dr. Eva Fabricius, J. C. 1801. Systema eleutheratorum, secundum ordines, Sprecher, Naturhistorisches Museum, Basel, Mr. Ole genera, species, adietis synonimis, locis, obsevationibus, descrp- tionibus. 2, Kiel. Martin, Zoological Museum, Copenhagen, Prof. Heikki Ferrer, J. and G. Moragues. 1998. Contribution à la connais Sance Hippa, Kjell Arne Johanson, Bert Gustafsson and Bert des Ténébrionides neotropicaux. Le genre Phymatestes Pascoe. Viklund, Naturhistorika riksmuseet, Stockholm to G. 1re Partie. Bulletin de la Société entomologique de France, 103 Curletti, Museo Civico Storia naturale, Carmagnola, and (4): 373–392. to Lukas Cizek and Pavel Prúdek, Faculty of Biological Ferrer, J. and G. Moragues. 2000. Contribution à l´étude des Ténébrioni- des neotropicaux. Le genre Phymatestes Pascoe, 2e Partie. Bulletin Sciences, University of South Bohemia, Czech Republic, de la Société entomologique de France, 105 (2): 113–123. for communication of material. The study was finan- Ferrer J., and G. Moragues. 2001. Description dúne nouvelle espèce cially supported by the Norwegian Research Council guyanaise du genre Phymatestes Pascoe (Col. Tenebrionidae). and the Norwegian Institute for Nature Research. Bulletin de la Société entomologique de France, 106 (1): 85–97. Ferrer, J., Barclay, M.V.L. and G. E. Hancock. 2004. Discovery of lost J.C. Fabricius (1775) and Olivier (1795) types of Coleoptera in the Hunterian Museum, University of Glasgow, Scotland. Annales References Zoologici, 54 (4): 775–781. Freude, H. 1967. Revision der Epitragini (Col. Tenebrionidae). Aalbu, R. and C. A. Triplehorn. 1985. Redefinition of the Opatrini Entomologische Arbeite Museum Frey, 18: 137–307. tribes in North America with notes on some apterous gen- Freude, H. 1968. Revision der Epitragini 2, (Col. Tenebrionidae). era (Coleoptera: Coleoptera, Tenebrionidae). The Coleopterist’s Entomologische Arbeite Museum Frey, 19: 32–143. Bulletin, 39 (3): 272–280. Gebien, H. 1919. Monographie der südamerikanischen Camarien Ardoin, P. 1959. Le genre Enicmosoma Gebien. Bulletin de lÁcadémie (Coleoptera Heterom.) nebst einer Übersicht über de indischen Malgache. Tananarive, 35 (1957): 60–77. Gattungen der Camariinen. Archiv für Naturgeschichte A, 83 Ardoin, P. 1961. Contributions à l´étude des Ténébrionides mal- (3): 25–167. gaches. Deux nouveaux genres dÁdeliini malgaches. Bulletin de Gebien, H. 1928. Über einige gruppen amerikanischer Tenebrioniden lÁcadémie Malgache. Tananarive, 37 (1959): 31–38. (Col. Heter.). Teil I. Stettiner Entomologische Zeitung, 89 (1): Ardoin, P. 1977. Contribution à l´étude des espèces americaines du 97–164, 167–234. genre Doliema Pascoe. (Coleoptera). Annales de la société ento- Gebien, H. 1937. Katalog der Tenebrioniden (Col. Heteromera). Teil mologique de France, 13: 1–20. I. Pubbl. Mus. Ent. Pietro Rossi Duino, 2: 505–883. NEW SPECIES OF DARKLING BEETLES FROM CENTRAL AMERICA 661

Gebien, H. 1938. Katalog der Tenebrioniden, Teil II, Mitteilungen der cations for estimates of global species richness. Conservation Münchner Entomologischen Gesällschaft, 28: 49–80, 283–314, Biology, 27: 283–296. 397–428. Ødegaard, F. 2003. Species richness, taxonomic composition and host Gebien, H. 1939. Katalog der Tenebrioniden, Mitteilungen der specificity of phytophagous beetles in a dry forest in Panama. Münchner Entomologischen Gesällschaft, 29: 443–474, 739–770. In. Arthropods of Tropical Forests. spatio-temporal dynamics Gebien, H. 1940. Katalog der Tenebrioniden, Mitteilungen der and resource use in the canopy. Eds. Basset, Y., Novotny, V. and Münchner Entomologischen Gesällschaft, 30: 405–436, 755–786, Kitching, R. L. (eds.). Cambridge University Press: 220–239. 1061–1090. Ødegaard, F. Host specificity, alpha- and beta-diversity of phytopha- Kaszab, Z. 1938. Die Arten der Gattungen Cnemeplatia Costa und gous beetles in two tropical forests in Panama. Biodiversity and Psilachnopus Reitter. Entomologisk Tidskrift, 59 (1–2): 77–83. Conservation, 15: (in press). Kaszab, Z. 1977. Neue Tenebrioniden (Coleoptera) von den Galapagos Papp, C. S. 1961. Checklist of Tenebrionidae of America north of the und den Antillen, sowie aus Südamerika und Südasien. Annales Panama canal. Opuscula entomologica, 26: (1–2): 97–140. Historico Naturalis Musei Nationalis Hungarici, 69: 117–130. Pascoe, F. 1871. Notes on Coleoptera with description of new genera Kulzer, H. 1949. Drei neue Tenebrionid-Arten aus Afrika und and species. Part I. Annals and Magasin of Natural History, 8 (4): Südamerika. Coleoptera. Mitteilungen der Münchner 345–361, pl. XIV. Entomologischen Gesällschaften. 35–39: 302–306. Philips, T. K., Steiner, W. E. Jr, and C. A. Triplehorn. 1998. A new Kulzer, H. 1961. Neue tenebrioniden aus Südamerika. (Col.): Entomo- species of Central American Platydema Laporte and Brulle logischen Arbeiten aus dem Museum G. Frey, 12 (1): 205–235. (Coleoptera, Tenebrionidae) recently established in Florida. The Kulzer, H. 1962. Neue Tenebrioniden aus Südamerika. Entomologischen Coleopterist’s Bulletin, 52 (3): 291–296. Arbeiten aus dem Museum G. Frey, 13 (1): 79–100. Pic, M. 1918. Nouveautés diverses. Mélanges entomologiques, 27: Kulzer, H. 1964. Über neue Tenebrionidenarten: (Col.) Entomo- 10–12. logischen Arbeiten aus dem Museum G. Frey, 15 (1): 221–276. Pic, M. 1921. Nouveautés diverses. Mélanges Exotico-entomologiques, Laporte de Castelnau, F. L. and G. A. Grullé. 1831. Monographia du 34: 22–29. genre Diaperis, Annales des Sciences Naturelles, 23: 325–410. Pic, M. 1935. Nouveautés diverses. Mélanges Exotico-entomologiques, Marcuzzi, G. 1953. Contributo a la conoscenza dei Tenebrionidi ame- 66: 1–36. ricani. 5. Nuove specie del genero Rhypasma, Atti Museo Civico Spillman, T. 1972, Uloma Dejean, Proposed designation of a type di Storia Naturale di Trieste, 19 (2): 73–86. species under the plenary powers. Z.N. (S) 1979. Bulletin of Marcuzzi, G. 1991. New species of Cyrtosoma Perty (Coleoptera, Zoological Nomenclature, 29: 32–34. Tenebrionidae) from the tropical region. Elytron, 5: 235–252. Spillman, T. 1973. Nomenclatural problems in six genera of Marcuzzi, G. 1994. New species of Tenebrionid beetles (Coleoptera, Tenebrionidae. Proceedings of the Entomological Society of Heteromera) from South America. Tropical Zoology, 7: 109–120. Washington. 75: 39–44. Mérkl, O. and J. M. Maes. 1991. Catálogo de los Tenebrionoidea Spinola, M. (Marchese de): 1844. Essai sur les Clérites, 2, p. 116. (Coleóptera) de Nicaragua. Revista Nicaraguense de Entomología. Sturm, J. 1826. Katalog meiner Insekt-Sammlung, pars 1, 207 pp. 17: 19–62. Triplehorn, C. A. 1965. Revision of Diaperini north of Mexico, Mäklin, F.W. 1884. Monographie der gattung Strongylium Kirby, with notes on extrelimital species (Coleoptera, Tenebrionidae). Lacordaire und der damit zünarchst werwandten Formen. Proceedings of the United States National Museum, Smithsonian Helsingfors, 517 pp., 4. pl. Institution, 117: 349–458. Olivier, A. G. 1795. Entomologia. Histoire Naturelle des Insectes avec Triplehorn, C. A. 1969. A synopsis of the genus Apsida with descrip- leurs caractères génériques et spécifiques, leur synonymie et leur tion of a new species (Coleoptera Tenebrionidae). Annales of the figure enluminée, Coleoptères, 1, .(imp. Baoudouin, Paris, with- Entomological Society of America, 63 (2): 567–572. out page numbers). Triplehorn, C. A. 1972. A revision of the genus Zopherus of the world. Ødegaard, F. 2000a. The relative importance of trees versus lianas as Smitsonian Contribution. Zool. 108: 1–24. hosts for phytophagous beetles (Coleoptera) in tropical forests. Triplehorn, C. A. and L. E. Watrous. 1979. A synopsis of the Journal of Biogeography. 27: 283–296. genus Phaleria in the United States and Baja California. The Ødegaard, F. 2000b. How many species of Arthropods? Erwin´s Coleopterist’s Bulletin, 33: 275–295. estimate revised. Biological Journal of the Linnean Society of Triplehorn, C. A. 1998. A revision of the genus Liodema Horn London, 71: 583–597. (Coleoptera, Tenebrionidae) with description of a new species Ødegaard, F., Diserud, O. H. and K. Aagaard. 2000. The magnitude from Costa Rica and Panama. Proceedings of the Entomological of local host specificity for phytophagous insects and its impli- Society of Washington, 100 (2): 324–330.

Received: June 15, 2005 Accepted: November 3, 2005